Skip to main content

Full text of "Palaeontographica Americana."

See other formats 


Р з... > 
N, EE ч r че у T + 


Е 
 PFaleontographica 
Americana - 


Fruits and Seeds 
of the 
Middle Eocene Nut Beds Flora, 


Clarno Formation, Oregon 


by 


Steven R. Manchester 


PALEONTOLOGICAL RESEARCH INSTITUTION 


Officers 
PRESIDENE Se oe Bork EL DUMP ee JOHN C. STEINMETZ 
NIT PRESIDENT $99 ао RICHARD E. PETIT 
ЕСЕРШ & 0 een re E n ЫЕ? HENRY W. THEISEN 
йок ап ошку ne ee ANNIKA FARRELL 
DIRECTORA. ee ee eg WARREN D. ALLMON 
Trustees 
R. TUCKER ABBOTT (to 6/30/96) PETER MCLAUGHLIN (to 6/30/95) 
BRUCE M. BELL (to 6/30/96) SAMUEL T. PEES (to 6/30/95) 
CARLTON E. ВвЕтт (to 6/30/95) RICHARD E. PETIT (to 6/30/96) 
WILLIAM L. CREPET (to 6/30/97) GARY ROSENBERG (to 6/30/96) 
J. THOMAS DUTRO, JR. (to 6/30/96) CONSTANCE М. SOJA (to 6/30/97) 
ANTON J. EGNER (to 6/30/97) JAMES E. SORAUF (to 6/30/97) 
ANNIKA FARRELL (to 6/30/95) JOHN STEINMETZ (to 6/30/97) 
HARRY G. LEE (to 6/30/97) SUSAN B. STEPHENS (to 6/30/96) 
ROBERT М. LINSLEY (to 6/30/95) HENRY W. THEISEN (to 6/30/95) 
Amy К. McCuNE (to 6/30/97) THOMAS E. WHITELEY (to 6/30/97) 


Trustees Emeritus 


HARRY А. LEFFINGWELL RAYMOND VAN HOUTTE 
WILLIAM P. S. VENTRESS 


BULLETINS OF AMERICAN PALEONTOLOGY 


and 


PALAEONTOGRAPHICA AMERICANA 


WARREN DSATLEMON o уу een EDITOR 


Reviewers for this issue 


BRUCE Н. TIFFNEY ELSE MARIE FRIIS 


A list of titles in both series, and available numbers and volumes may be 
had on request. Volumes 1—23 of Bulletins of American Paleontology have been 
reprinted by Kraus Reprint Corporation, Route 100, Millwood, New York 10546 
USA. Volume 1 of Palaeontographica Americana has been reprinted by Johnson 
Reprint Corporation, 111 Fifth Ave., New York, NY 10003 USA. 


Subscriptions to Bulletins of American Paleontology may be started at any 
time, by volume or year. Current price is US $50.00 per volume. Numbers of 
Palaeontographica Americana are priced individually, and are invoiced separately 
on request. 


for additional information, write or call: 


Paleontological Research Institution 
1259 Trumansburg Road 
Ithaca, NY 14850 USA 

(607) 273-6623 
FAX (607) 273-6620 


©) This paper meets the requirements of ANSI/NISO 239.48-1992 (Permanence of Paper). 


- Paleontographica 


_ Americana 


Begun in 1916 


| 
| NUMBER 58 AUGUST 24, 1994 
| 


Fruits and Seeds 
| of the 
Middle Eocene Nut Beds Flora, 


Clarno Formation, Oregon 


by 


Steven R. Manchester 


Paleontological Research Institution 
1259 Trumansburg Road 
Ithaca, New York, 14850 U.S.A. 


ISSN 0078-8546 
ISBN 0-87710-435-2 
Library of Congress Card Number: 94-65491 


Printed in the United States of America 
Allen Press, Inc. 
Lawrence, KS 66044, U.S.A. 


| 
| 


CONTENTS 
Page 
A ee een ee a о е ое ee. Den Inc o г: 
чо ома (еко ти (ери Ни Б с ке а Te x cd E Е а 7 
ОСОО е ын сокет тена ЕСЕК К ек eco До не оо бойун киш каек en Se 8 
сорар аео Sein ы A аве скн ы at AEC mh хел Red Гут ue qM т ЖЕТ 9 
OE реве DMN ы Вир aN ы ML и ди ы кю ы О ыры UM С ааа en 12 
О О Раоа ЈАНО а А И Кеде ee a eas ei cU rm ERO eh 13 
MOU BSSOfaBLESETVaON е со СЕ M а c c M M iT ы mb 13 
Маи М СТВО И ee уы са E ыу bade ои 14 
СЕЗА ыст уы Аш ык mos TUS du win vad а СК кы зк ке ы жо en A 15 
Sy От DN МВ Sab ӨӨ en ee ose c ME M Mc oce cm А 15 
¡STO WMA TURO dais por Бо НУ ама а не канау лж М ке Е ее cud Crew chute) тул e г: Ж 19 
Саа Ө ИЕШЕ ШӘ сокет и за earns, С ЕЕ mS s dae. 22 
BIOECOETADLICKEONSIUSLHLIONSE ео 23 
(ВО РАНО Ко Бети Бо Ши ӨТӨН sac ee 23 
ООШ ОШКОШ ШИК о ТЕ ШАН С а ин чс ез eu rec ees Е а ИЕ oy c у ae у ды 26 
OLSON OSY a БЕ ЛООНУ узу Иол. wy оса а и Ер у Ай 27 
Ore ыссы ыы MIT ee een 28 
Шека DOOR SR а ee ee АК 28 
DGHSUTCIDOnisPe NONU c ue D M e era ins Cu mU MD MO bd Cue QR DRE A m AME 30 
SDDLENIAUONSE RS cec боза ы cte ed ee ola ey Gee ME он се claim E me 30 
Conifers 
Пи а Mme AX. ee ccu d e C tas KR E EMEN ENSEM оа 30 
IHE EAE mE es c die M ed uec etu P а dm 30 
ШОК СОЕ КН amc cc UM NECEM UC eL SE CE c MR EE c Ей 31 
И промо Mp жар. P cc ur I A скы m CL NEL Mara ed cs tutior itt 31 
WOMAN ВАРНЕ REESE, c p Md pM E I ове mu EM mM e rdi 
ПРОРОК УНЕ UE de o ctu ie exi ee en tds esa A E E ede esed rc plaid 32 
Angiosperms 
ООА СОВЕ M Mee t eU Кы HL UU M I MC m d Muti Ja 33 
A лы без син оао D e Чу К е С O cU Werde Lie s СИ ын o, 33 
Е ЕО О ИИИ ИАА 34 
Zr d а ан Ма dM x M P I MA ek 34 
AL Та ел ОМА ИЕ M itr Cha wan I cec E NU MUN S AS EU 35 
ПАЗИТИ UU ep кө cR D que rcc M eL o iM ae TN 35 
MU ui uum. a ep M ON uu c Ка e mM er Mr mae ы A 35 
END IE Sem ee ee ee ee e E 37 
EIN ON ОВИ TL) ОЕ e ee Rc а и c. = 37 
TAN ACEAE AN N tr na OLE и лане к ОВ RO LL ее е пе tc а О окови 38 
ОРИ ИЕ EONA Син К ee Mee N Rad dI LU edt E EE, НИ 38 
BCT ACU AORN а LE ои MM Ware па о и иза cc ко | = M 39 
ISOTOTSDOUTIORDOD NONAS nih cain шешек ен RE CI к ети wah ede wean car Mees И Ne ыа E реа е ы у 39 
ООС ОИ тама ре м беседа е и на е Le UR cm E M ee QI МЫ ы ы и tU 39 
ШКЕАК Н единна ИИ INN fa дан erc тл O ко е CT ae 40 
IA ү ы у к е NU ВИ Gu soa ы кыч ER e eet АК темя 40 
SOT ACen И, Do MID уны uc сы c c E S Lu 41 
[ОП КИЕК PAIL ARE Dd RM СИВ cues E эу E кта ме у КЫ о AN a АНУ бт ы г C er. . НИ 41 
дум роле EE ы е qua M Би ЕН а тез reU UM и к лы ee ey I 42 
Ди но OE aie акны Dy Gap ea RN tone uud c шерт x M uuu cM LE P E 43 
КОЧО АИ ОЈ MERE cic орат Г л м ме ым С оо чек NM ee тосе сус E EN 44 
ЕТО они ЖИК POCO ee ee direi. uade: 44 
D ta ра БР Еа БЕ ма 45 
а ет OR лы d E у ccr x nen 46 
ЧО О ОЮУ ah E Ок A kno а HI ett 46 
(Ори Ке Дуа ED О ME c ен oe ba Е en ис о PP dc d cM E M лоо Лиао NEHME NUT 47 
ШЕ ШКЕ Ин ее oltre оо Co о ы c qu c Муш по P mI 47 
A АЕ аа ро ену De dra MM Hu up nc ME MT 48 
ПУТА БИ ДОЊА oc эз P LN pd pm I M I NM S CE E 49 
КОЛО ESE О КОХ КИШКЕ А M Miu 49 
ШОЧ КШ КЕДЕ ee REEL Nd C Lo лл RN i. qu. e ri d ER CU 50 


КОЛЛ АР em Me dv ge M eMe x ep M st TN л ое г ee a 50 


T d Ty a кк Кы cc у озук ИННИ О кн E DNE ы О л, 
И анас РОТИ di OIE FS OIE RE FREESE у nit н 
oe рол NN ух сим у ее л и у нил c MS nee RM 


URN Оос OR л к и к ee ee A 
И е лы кор е р сы сс a RE 
P ЖО шс аа ко ы с т ко сс MA tee И 
Жк Сол со Lo ee T pue ре Ma Е I QUEEN ei Aa ЧИН Vicus 
Dp xeu ccce esL VEL m 
N ръста рапа ш у o о ныды ENANA 


ee o ER CL Du ОО ERA 
PET lu m s I Um ec uU dene MEER AULAM 


КОИ ен or nion a ro Duk О ua epi pneu Ex NE 
NNNM. eoo е су с ann ат oe E a ade ка КАЛИ ee ы 
A a E a M О ООО АО E 
E RM S ксы жо S c I UI d UE 
(oor c M TP LIN MI IE vr кес cuu S MD MM UM ME 


ciusdem ecu К узо ee E eM M 
КИМЕ на tu uu СО EA ОА d I D M АЕ 


| Оа аб o deor uote ee death, cee e Ed К шы Net UU е ни EN EEE IE A a вай 90 
| FAD HGMGNTH ем ne канам Ee hatte EN Ce at ee Е КУМ оа cd. da erar 90 
| WEISE ee ae c cn Macte og E СИ ы NEUE ee 91 
d ШЕЙ dela нс E E А E E ee I M E d 92 
| ECA CLOSDERIN т М ж E E N ee Н РА м 92 
ОВА a E И M tU MR CL cu О DUE у кн A EUM күзү т iE 

| LAINDEIOCISSUS о arith eater пи ин E ЛОТ КККК EN ЛУ т о itn coe на е add 93 
| TAS RA NS ЕЕ СУ ee NE a UN ee er Re CN om m е 94 
AA EIER пена dees er eL ee А КАФ dud d 95 

| а BE. они о ооа В а у E 95 
есу ере е ен ИНО 96 

| PS LODO ПАВЕ Оа cR I MUR utum Tc uuu c t ay RE 96 
PATON ОЛЕ А ON A aa E Кол, Жек T A MEE t t Lc I а ur у ет 97 

3 AI ee en ED CET D I D UU MM tee cd more A ar 97 
| AXINOSDENMAEBONENON ENE АЯ SE Ea Ку eL MEE УЕЛ S ЕГ ea MA Кук ME MUN ТА SA 97 
ОПСТАО DO МАЊА e NE NL СЕН АА E RT EU Е ОКА, жо СС OP AO 98 

| (БОРИСА е EEMO T Oe IX MM TOM NE M Ice Keen ctm ДРЕ ТС а CUM 98 
\ (КИЛЕШӘ ЕШ nove UR rM SI РЕА Se NE ou И Р ON PULLUM пе се ар 98 
| ЖОЮП е ИОН ENERO Vitex е NO ЕНУ куз ORUM SC DU. ue cM EU ICI TERES DUET Ld. ORA 99 
DULOCALPUS ROME ILO Va er eee ed АИ ИО MM MUSEI MEE PC ME eI UR PP аа 99 

| BEUNO CERU EORNA E E И Cc TO: OS ON МОР АТИ Mu M ЕЕС КИТЕ В AUS СР аре: AN 99 
| LTD TALL CODOS E EU ЕТ ТИЛЕ Е A ER н оа ем ад 100 
EFA SATILES Bele DOV ее РЕНЕ Са Н, IPM С e A ee У А a E E 100 

| ФгоБийсавртринвейе О ИИ ТОИ NR АБА OM PEU M EAE Iti PM 101 
ПЕХАР РО НО A m T Л НИСЕ НИ E NI UM NUMINUM e eee 101 
JOCJOHGSIQCECBUTIOVIR NE eee BS НИНИ REESE LUE NEMUS E ON OAM МОЛТ ee EE 102 
LighicanpuSpenenov dme qe LR ET Ee ete SS PIDEN А аа. 102 
LiehigioD аа MON NS NI CHRIS PUT сае RE MET СИ EU ELEME UE MEUSE ICM 102 

VEGA HON (helo ON ON DONA EE au ase SHENG Т TU e MUN E Rm M CI RN MEE I UM NEC E 103 

| ОЛОР И ӨЛӨ ОИ A за ЭШИ ОНЫ NP IS ERD c ЖЕЛ О А ee КЕЧ 103 
| INEDAFOSCMPN genova ee НА И 103 
ОПУВИТРУШТИ ВОН ИО УЛ РАНИ N O A S 104 

| PAStePHAGKTGISEN ENO Ve cU N AR TEN A ED MOX IEEE реа Пр маже DICE 104 
РИО БЕЛ BEN PHONE ОН Ыы win КЛИШЕ EON Е cite. A eee cM СЕЕ. US A А MS 104 

| Раббы ох н а Уа а а а А И cO ОРЕГОН none mn 105 
| POUOSTOSDENINA:DENEN ONE АНЕ E ET КК А odes ep NE Feder eed ОКНО ЮКО A 105 
| POL) STANDISH ONE ER трона TN TE А Em Пи AAA ANOS NEE 106 
i Erun DO POMELO TET Ку Vd e E A. TA O A ж AR ER 106 
ТЕГОВЕ CEN NOVERIT rM, AOS МЫ ТЬ AAA ERA ER cT TAA 106 

| ШИРЕ И РОВ пот Pee ee ee A LOST Пи au ШЕЕ 107 
Уа ДЕА епа торта ИТЕ LOUP AE Aceh И Н О ИТО E O PA О E O а. 107 

| ОПИСИ ОЛОХ Е МАТИ RE Виан ара cU earl ААИ Ра а E Ба 107 
: АОИ Чат ато Жы II К Om Cok ia все RRM т TEN 108 
| SS ЕПОРАССИ ТРИ И Сево У AL О И qoe mox ios NIMES 108 
DERLATIENGEIEBENE ONE ee а steilen 109 
СПРИТНО ООУ N КОС атта oe MUR 109 

MS DUGETOSPEHING Belle NOV EA A A O E r 110 

| ISDREROSDELINGEBENNOV a УРААК УУ TIE D V И ee le ee а ПАДА НЕ I EN 110 
| Saena ananena e А o uS oe een. ee era ee 111 
реА ОУ A АДЫР S AE Udo рз жөө Sae E аб КЫ шыбы рыб ыан ааа cus ли uisa RN 111 
ПОПИО ве ОМА ch tak teeta LC rer ЛЕКТЕ ка re i qM M onn 112 

Ши тесари реши ON АЛ И ЛУ е ери те RE ЛЕ УУУ ЫКЫ EAS ANCA CUM LUE PORA AAT DET T e Сулк PT RN 112 
LEONOV LE POMC ОМ АУЕ УГЕ Т ЫЕ ТУУЛ УЛТ УТ ШУ УУУ ЫТ лел пра а ТУМО NEIL ИЕ Co BR PEE 112 

| IDAS TC MSE CIATION AME ENTER, eRe e ЫЛ ER DL. AS, EOS BERO IE wey N тоа A 113 
LTO OSCOPNG ДӨШҮ A Е ES ue OO Ioco ee ee A rn BR 113 

| TES EMITIR TE ES DI OS ORTOS TOTO CLIQ л E E ВИИИ ао RT пе MN 114 
TA NADIE MAME M M D M Me aue QN i и 114 
ТШ AA О УО PIA T S UE E NNI CE e AS 115 

СПОРЕ ТИТ ВОНО OR ar OI Ууу QNO AL, МЕ MRAM UL Т LEER ERN RH 115 

REAT TNE NONN TE tre AM Fie ОКК S EMI ct EDO EN, EN NER adi 116 

| (РОДИ И ЛЕЛЕРИ ОНЕШ ШЕР А СИ ое iiu. Beate learn, 116 
(Селма HAAS oF у EROR Жил Аск Ко с а Ne АЫ е ELI NUR M о ECL. 117 


TAAA ЕР c лла ы сие T О РМ Ж AN Ci еа M RE UE ta кх ed 117 


Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 
Carpolithus sp. 


References Cited 


Index 


Text-figure 
. Geologic map and index map showing location of the Nut Вейѕ.............................. n nn 


LIST OF ILLUSTRATIONS 


ЕК E E E Не оно ШУ se eus er ee Моо О ШО ОО uae л et 
ци опело ae Эз UO EIC UM cM IC MIR Eq 
ЕЕ ОВОО ше чк к y THE АН ек AUN TR cR hog DA CE Gee OO E A ы А 
c Taxonomie status об Claro fruit and всей репета esi cov e b rrr retrum о иска ааа d rg t rn neue toe tes 
= Nut Beds genera ти не Eocene об Богоро oe aye che стани td Ag peur en usua 
. Present geographic distribution of extant Nut Beds genera ......................... emen 
Es cH IP Hd LC c E M LAU ДИ е SS qr d Ne eh ATE IP бнр Tte sud. 
И ОА НА te Lo I M аа ен cs eger E а Т cue uL E 
BANCOS OO C D T RAS eth UU A UTEM C M T 


SE ID DM C dM URL Mo Vm DIL M A ОО ОО ОО A Mo EMI 5 
POR ИРИ LL ОЛО Е IT MS Eod serve iue огл edu cilc Dua лы 


N ee ORO ON QS ONE E I ERN an TIN UU 
ЕРА = UT ECCO bocce tn INN cO eL RM E T 
. Emmenopterys . 


TWO TU cc e TER IR UO e LU WP CIS er E 


Table 


1 
2 
3. 
4 
а 


= ООО о Acer and: Deviacer samata oer coir. БИ Rene кома е вота Oh Roi Mew ин б ee э жүзө жыды” 
. Ferrignocarpus . 
. Fimbrialata ... 
. Pteronepelys ... 
. Quintacava .... 
. Scaphicarpium . 
. Sphenosperma . 
. Triplascapha... 


LIST OF TABLES 


. Comprehensive list of fruit and seed taxa from the Clarno Nut Beds assemblage indicating number of specimens observed ..... 
. Present habitat and vegetational distribution of Nut Beds genera ............................................... шы 


Tasa ЭНСЕП WILD European ОСО MOTAS аи а И Y ee e V Qe De Y E eee niu 5 | ка ае 


. Present geographic distribution of extant genera represented in the Nut Beds Нога... ... 0.0.50 e seen 6 
. Systematic list of Nut Beds genera with known modern familial affinities . . ...... eee nn 


FRUITS AND SEEDS OF THE MIDDLE EOCENE NUT BEDS FLORA, 
CLARNO FORMATION, OREGON 


\ 
| STEVEN К. MANCHESTER 


Florida Museum of Natural History 
| University of Florida 
| Gainesville, FL 32611 


ABSTRACT 


Permineralized fruits and seeds from the Nut Beds flora of the Clarno Formation, north-central Oregon, are described and 
\ illustrated, providing the first comprehensive treatment of a North American fossil fruit and seed assemblage and facilitating 
| comparison with European Tertiary fruit and seed assemblages. Newly obtained radiometric dates, along with vertebrate cor- 
| relations, indicate а Middle Eocene age of about 44 million years for the assemblage. Based upon examination of approximately 
| 20,000 specimens, 145 genera and 173 species аге recognized, providing much new data for systematic, evolutionary, biogeo- 
graphic and paleoecologic interpretation. Seventy-five genera and 102 species have been identified to 35 living families; 70 genera 
| and 71 species represent form genera of unknown familial affinity and may belong both to extinct genera and to extant genera 
whose affinities may come to light with future work. 
Among the 102 species attributed to extant families, 58% belong to extant genera, 30% are extinct genera, and 22% are 
| stereotype genera (identical in the characters preserved to more than one modern genus). Trees апа lianas are well represented, 
nearly to the exclusion of herbs. The only herbaceous angiosperm identified is Ensete (Musaceae). The proportion of the identified 
taxa that represent lianas or climbers is high (43% of the 69 species for which growth form may be inferred) including 14 species 
of Menispermaceae, seven of Icacinaceae and six of Vitaceae. The disseminules range in size from about 0.5 mm to 85 mm; 
about 15 are winged and suitable for wind-dispersal, but most are medium to large in size and represent drupes and berries 
adaptive for biotic dispersal. 
| Among Ше extant genera determined, some are exclusively temperate in distribution today (e.g., Taxus, Emmenopterys, 
| Parthenocissus), while others are exclusively paratropical to tropical (e.g., Ensete, Mastixia, Iodes, Ampelocissus, and, among 
the vertebrate fossils, a crocodilian). Many of the genera are distributed today in both temperate and tropical areas. Despite the 
| оссштепсе of many taxa that are temperate today, the presence of Musaceae, Palmae and other taxa that typically are frost- 
intolerant today, combined with the high diversity of lianas, is taken as an indication that the climate was comparable to that 
| which supports paratropical rainforest vegetation today. The flora shares 24% of its genera with the Early to Middle Eocene flora 
of western Europe, indicating that one or more land connections were viable during or prior to the Middle Eocene. Among extant 
genera represented in the Nut Beds assemblage, the greatest similarity is with the extant flora of eastern Asia. The identified 
families are, among the conifers: Taxaceae (3 genera), Pinaceae; among the angiosperms: Actinidiaceae, Alangiaceae, Anacar- 
diaceae (2 genera), Annonaceae, Araliaceae, Betulaceae (2 genera), Cornaceae (6 genera), Fagaceae (2 genera), Flacourtiaceae, 
| Hamamelidaceae, Hydrangeaceae, Icacinaceae, Juglandaceae (4 genera), Lauraceae (3 or more genera), Leguminosae, Lythraceae, 
Magnoliaceae, Menispermaceae, (13 genera), Musaceae, Palmae, Platanaceae (3 genera), Rosaceae, Rubiaceae, Sabiaceae (2 
| genera), Sapindaceae (2 genera), Sapotaceae, Schisandraceae, Staphyleaceae, Symplocaceae, Theaceae, Ulmaceae (4 genera), and 
Vitaceae (4 genera). 


ACKNOWLEDGEMENTS Museum of Science and Industry, including John M. 
| Armentrout, Bruce Hansen, Michael С. Houck, апа 
| This work is dedicated to the memory of Thomas Joseph Jones, III, and members of the Oregon Agate 


J. Bones (Text-fig. 1), who collected the majority of 
Specimens upon which this investigation is based and 
whose enthusiasm helped to kindle my own interest 
in paleobotany as a high school student. Many others 
assisted me in collecting at the locality as participants 
in paleobotanical programs sponsored by the Oregon 
Museum of Science and Industry during the summers 
of 1972 to 1989, including Alexander Atkins, Scott 
Blanchard, Kris Goertz, Тап Gordon, С. Bruce Hanson, 
Carol Hardman, Elizabeth Harding, Kathy Harvey, 
Joyce Lenz, Maureen Muldoon, Duane Olson, Jerome 
McFadden, Ellen Pasternack, John Ries, J osephine 
Spitzer, Michele Vowell, Eric and Mike Weinstein and 
Others. I thank former and present staff of the Oregon 


and Mineral Society, including Gar Hurley, William 
and Gertrude Hall, for their support of student field 
programs in Clarno paleobotany. 

Special thanks are due to Carrie L. Roose, for in- 
valuable help in sorting, measuring and cataloguing 
specimens, assistance with initial taxonomic analyses 
and in preparing the geologic map and preliminary 
diagrams. C. Bruce Hanson provided help in inter- 
preting the local geology. Wendy Zomlefer assisted with 
preparation of the text-figure illustrations. David 
Dilcher provided access to equipment and facilities as 
well as helpful encouragement throughout this re- 
search. N. Gary Lane kindly provided access to a Mi- 
croslice II annular diamond saw for sectioning the seeds. 


8 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Text-figure 1.— Thomas J. Bones, photographed in 1977 at an 
excavation on the top of Face 3. Mr. Bones collected here for about 
40 years, and provided the massive collections upon which much 
of this monograph is based. 


Advice in the preparation of latex and silicone casts 
was given by William Simpson and Russell McCarty. 
Help with identifications was provided by Margaret 
Collinson, Robert Kaul, Peter Endress, Paul Grote, 
Dieter Mai, Charles N. Miller, Richard A. Scott, and 
Bruce Н. Tiffney. Grammatical advice on the construc- 
tion of new Latin binomials was provided by Benoit 
LePage and Christine Kampny. 

Access to paleobotanical collections was provided 
by Scott Wing, Francis Hueber, and James P. Ferrigno 
(U.S. National Museum, Washington, D.C.); Howard 
E. Schorn (Museum of Paleontology, University of 
California, Berkeley), Wesley Wehr (Burke Memorial 
Washington State Museum, University of Washington, 
Seattle), Charles Beck and Robyn Burnham (Museum 
of Paleontology, University of Michigan), Andrew 
Knoll (Harvard University), Leo Hickey (Peabody 
Museum of Natural History, Yale University, New 
Haven), Chris Hill and Cedric Shute (Natural History 
Museum, London, England), Christiane Blanc (Mu- 
séum National d’Histoire Naturelle, Paris, France), 
Friedemann Schaarschmidt (Senckenberg Museum, 


Frankfurt, Germany). Access to extant fruit and seed 
collections was provided by Charles Gunn (National 
Seed Herbarium), Peter Stevens (Harvard University 
Herbaria), Norris Williams and Kent Perkins (Uni- 
versity of Florida Herbarium). Helpful comments on 
the manuscript were provided by David Dilcher and 
Herbert Meyer. I also want to thank the reviewers, 
Bruce Н. Tiffney and Else Marie Friis, for invaluable 
comments and suggestions. 

This research was supported in part by the following 
National Science Foundation grants: DEB 81-11-89, 
EAR 8707523, EAR 8904234, BSR 9007495, EAR 
9322765. This publication represents work that was 
initiated with the facilities and support of Indiana Uni- 
versity and completed at the University of Florida. 
Publication costs were met in part by funds from the 
Gatorade Account of the University of Florida. This 
publication is number 410 in the University of Florida 
Contributions to Paleobiology. 


INTRODUCTION 


A diverse assemblage of well-preserved Middle Eo- 
cene plant remains occurs in the type area of the Clarno 
Formation about 3 km east of the community of Clar- 
no, north-central Oregon, in a locality known as the 
Nut Beds. The site yields abundant fossil woods (Scott 
and Wheeler, 1982), leaves (Manchester, 1981), and 
exquisitely preserved fruits and seeds (Scott, 1954; 
Bones, 1979). Because fossil fruit and seed floras are 
relatively rare in North America, the Clarno Nut Beds 
flora provides an important link for paleobotanical 
comparisons with the Tertiary of Europe, where fruit 
and seed assemblages are abundant and relatively well 
studied. The Clarno flora often has been cited in phy- 
togeographic discussions because taxonomic similarity 
with European fruit and seed floras provides evidence 
of broad floral continuity between Europe and North 
America during the Eocene (Scott, 1954; Chandler, 
1964; Wolfe, 1972, 1975; Tiffney, 1985a, 1985b; Col- 
linson, 1988; Manchester, 1988). Only a small portion 
of the Clarno flora has been described, however, and 
the actual extent of generic and specific similarity be- 
tween the Eocene floras of North America and Europe 
has not been well documented. 

The paleobotanical and phytogeographic signifi- 
cance of the Clarno flora was first shown by Scott (1954) 
in a monograph detailing the morphology and rela- 
tionships of ten species from the Nut Beds locality. 
Scott’s work demonstrated the presence of genera with 
Old World tropical affinities in the Eocene of western 
North America, and documented several genera in 
common with the Eocene London Clay flora of En- 
gland. He also called attention to the presence of both 
extant and extinct genera, a finding somewhat at odds 
with Tertiary leaf studies of the time. Subsequently, 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 9 


an expanded taxonomic list of Clarno fruits and seeds 
was published (Scott, in Chandler, 1964, p. 58) in- 
cluding 32 additional genera from the locality. Until 
now, these additional taxa have not been formally de- 
scribed. Bones (1979) published an atlas depicting some 
of the diversity present in the Nut Beds fruit and seed 
assemblage, including a variety ofnew but undescribed 
species. Several additional fruit and seed taxa from the 
Nut Beds have been treated in separate accounts (Man- 
chester, 1986, 1987a, 1987b, 1988, 1989a; Manchester 
and Kress, 1993), but a comprehensive treatment of 
the flora has not been attempted previously. The Nut 
Beds fossils represent one of the warmest climatic in- 
tervals in the Tertiary of North America, a time when 
thermophilic vegetation was spread well into the north- 
ern latitudes (Wolfe and Poore, 1982; Wolfe, 1985). 

Because of the excellent details of morphology and 
anatomy preserved in silicified fruits and seeds from 
the Nut Beds flora, specimens from this assemblage 
provide the rare opportunity to investigate the internal 
structure of Eocene plant reproductive organs. Such 
specimens enable a well-informed assessment of tax- 
onomic affinities, and make it possible to test whether 
affinities inferred from external morphology are upheld 
by internal anatomy. The Nut Beds specimens thus 
provide insight into the structure and affinities of var- 
ious extinct and extant genera of fruits and seeds that 
are known only from molds or impressions at other 
fossil localities. 

This monograph is an attempt to provide a thorough 
taxonomic treatment of all known fruit and seed spe- 
cies from the Nut Beds flora, including taxa of unde- 
termined familial affinity as well as those which can 
be identified confidently to modern families and gen- 
era. The resulting data base: 1) provides new systematic 
data on plants present during the Middle Eocene in 
western North America; 2) contributes information on 
the evolutionary status of various gymnosperm and 
angiosperm taxa in the Middle Eocene of the northern 
hemisphere; 3) provides insight into the environment 
and climate under which the Nut Beds vertebrate fauna 
existed; 4) enables comparison with other Eocene fossil 
floras of the northern hemisphere and to modern flo- 
ristic regions to gain some understanding of timing and 
routes of phytogeographic exchange. 


GEOGRAPHIC AND GEOLOGIC SETTING 


The Clarno Formation is a terrestrial sequence of 
andesitic to basaltic lavas and intrusives, ash flows, 
volcanic mudflows, and tuffaceous sediments exposed 
along the Blue Mountain Anticline in north-central 
Oregon. The distribution, stratigraphy, age, structure, 
composition and source vents of the formation are 
reviewed by Walker and Robinson (1990). Petrologic 


and trace element studies suggest that the volcanism 
may have been in response to subduction zone mag- 
matism (Rogers and Novistsky-Evans, 1977; Noblett, 
1981). The formation is well known for its paleonto- 
logic importance and is the focus of the Clarno Unit 
of John Day Fossil Beds National Monument. Leaf 
impressions and silicified wood occur in tuffaceous 
strata in many areas of the Clarno Formation (Hergert, 
1961; Manchester, 1986, 1991). Fruits and seeds, which 
occasionally occur in the same strata as leaves, are 
usually preserved as impressions or compressions. 
However, the Nut Beds locality is unusual because it 
provides uncompressed, three-dimensionally pre- 
served, permineralized, fruits and seeds. 

The Nut Beds locality (UF loc. 225) is situated in 
the type area of the Clarno Formation, at 44°56'36” N 
lat., 120?25'34" W long. (SW 4, SE Ya, sec. 27, T 7 S, 
R 19 E), Wheeler County, Oregon (Text-fig. 2). It is a 
resistant, buff-colored, cliff-forming unit about 10 me- 
ters thick, exposed over an area of about one hectare 
along the western side of Hancock Canyon (Text-fig. 
3). The term “nut beds" was first applied by amateur 
fossil collectors in the 1940's because of the abundance 
of petrified walnuts and other fruits and seeds found 
there. Fossil plants occur throughout the vertical and 
horizontal extent of the exposure. The same locality is 
also significant for its vertebrate fauna, which consti- 
tutes the only known Middle Eocene terrestrial mam- 
malian fauna in the Pacific Northwest (Stirton, 1944; 
Hanson, 1973, p. 56 in Retallack, 1991). 

The complex geology of the local area surrounding 
the Middle Eocene Nut Beds locality, and stratigraphic 
relations with the Late Eocene/Early Oligocene Han- 
cock Quarry vertebrate locality 1 km to the north (Text- 
fig. 3), are currently under investigation (C. B. Hanson, 
pers. comm.; G. Retallack, pers. comm.). Hanson (pers. 
comm., 1992) distinguishes five unconformity-bound- 
ed units in the type area of the Clarno Formation. The 
Nut Beds deposit occurs low in the section, within the 
second unit, and predates an adjacent andesitic plug! 
which was a source for clasts found in subsequently 
deposited sediments of the Clarno Formation in its 
type area. The Hancock Quarry fauna (Mellet, 1969; 
Hanson, 1973, 1991) and flora (McKee, 1970) locality 
is situated in Hanson’s uppermost unit, an upper Clar- 
no valley-fill deposit containing clasts from the an- 
desitic plug. Retallack (pers. comm., 1993), however, 
reports finding clasts of the andesitic plug! in the Nut 
Beds sediment and considers the Nut Beds to correlate 
with the upper part of a sequence of lahars and tuffs 
exposed | to 2 km north and east in Hancock Canyon. 

The Nut Beds deposit is composed of tuffaceous silt- 


! New information available; see Notes added in proof, p. 200. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


IN 


SR 
A SS RS SA 
ANTÉLOPE 2 AR А 
oy 4 


GSSIL 


USN — ||| 
S \ A QUIET 
NA N A e i l 
D £ s А | 
cy po As | қ | 
ASHWOOD 47, 2» Хам | 
: 9. ? E TWICKENHA | 
‘ip, {ч M 
UIT Q 5. 
wy сас | 
) MUT | 
S N SE ern 
зе = | 
pto === | 
Y) хе S4 m. im) 
$ wr, — MITCHELL | 
(n 10 BETTER ai NN | 
V Se | я 
Ata euer | 
N тс AT 
i | 
Y [ RU , 
N | Plio/Pleistocene 
SS ||, П сее |) 
В |) 3 | 
S SE N abel 
~ ` ormation 
AN а жале Еосепе С!агпо 
9POST Е N RR Formation 
Ve N SEE Cretaceous marine | 
= f Sun ] sec 
ка SS vi 5 10 15 Mi 
у E 10 15 20 25Кт 


Text-figure 2.— Geologic map and index map (inset) of the John Day and Crooked River Basins, north-central Oregon, showing location 
of the Nut Beds (arrow) and other paleobotanical localities of the Clarno Formation. Localities: 1. Nut Beds, 2. Hancock Quarry, 3. Dry 
Hollow, 4. Left Hand Canyon, 5. Horseheaven, 6. White Cliffs, 7. Indian Rocks (classic Cherry Creek site), 8. Red Gap, 9. John Day Gulch, 
10. Ochocco Summit, 11. Doolittle Flat, 12. Alex Canyon, 13. Classic West Branch Creek locality, 14. Brummers Spring, 15. Sheep Rock 
Creek (Teater Road site). Geologic map adapted from Walker (1977). 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 11 


i s Se 


Text-figure 3. — Oblique, northward, aerial view overlooking the Middle Eocene Nut Beds locality, with exposure faces labeled 1 to 5. The 


SEN } = 


dark unvegetated strata overlying the Nut Beds are a sequence of paleosols locally called Red Hill (RH) that is capped by the basal ignimbrite 
of the John Day Formation. Other localities in the distance include the Upper Eocene Hancock Quarry (HQ) in the upper part of the Clarno 
Formation and the Oligocene Dugout Gulch leaf locality (DG) in the lower part of the John Day Formation. Iron Mountain, capped by 


Miocene Columbia River Basalts, forms the horizon. 


stones, sandstones and conglomerates thoroughly ce- 
mented by silica and calcium carbonate. The strati- 
graphic column at the Nut Beds is about ten meters 
thick, consisting of a lower half with alternating beds 
of siltstone and sandstone mostly 0.1-0.6 m in thick- 
ness, and an upper half with thicker beds of alternating 
siltstone, sandstone and andesitic conglomerate (Text- 
fig. 4). Clasts of the conglomerate are subangular to 
rounded, generally less than 4 cm in diameter, and are 
matrix-supported. Permineralized wood, fruits and 
seeds occur throughout the vertical section, in siltstone, 
sandstone and conglomerate layers, with the greatest 
concentrations being in the uppermost strata. The ver- 
tebrate fossils are confined to the middle and upper 
strata, often in conglomerate, and are usually preserved 
as disarticulated bones. Impressions of leaves occur at 
various levels within the deposit, but are most abun- 
dant and well-preserved in siltstones near the base of 
the section (Manchester, 1981). 


Abundant Equisetum stems and occasional fern rhi- 
zomes are found in growth position in some of the 
siltstone layers. In some parts of the deposit Equisetum 
stems may be traced vertically through 0.7 m of sed- 
iment (Retallack, 1981), indicating relatively rapid 
sediment accumulation (Scott, 1954) and shallow-wa- 
ter conditions. Predepositional abrasion is evident in 
some of the fossils (particularly fossil. woods in the 
upper part of the section) and suggests that the assem- 
blage includes transported as well as locally derived 
plant debris. Although well-preserved silicified wood 
is common, most pieces are small, predepositionally 
worn fragments; no standing stumps were confirmed. 

The Nut Beds deposit is dissected by slumps and 
erosion into five outcrop faces, which, for reference, 
have been designated from south to north as Faces 1- 
5 (Text-fig. 3). Most of the vertebrate remains were 
recovered from the middle and upper portions of Faces 
1 and 3. The fossil fruits and seeds described by Scott 


I2 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Text-figure 4.— Profile view of the stratigraphic section at Face 3, 
showing alternation of beds (siltstone and sandstone/conglomerate). 
The lowest exposed layers contain leaves as well as some of the 
fruits. The most productive horizon for silicified wood and fruits 
was found to be in the massive upper layers. 


(1954) and the Hyrachyus tooth described by Stirton 
(1944) were obtained from two beds situated in the 
gully between Faces 2 and 3. Thomas Bones collected 
fruits and seeds throughout the deposit, but found the 
richest horizons in excavations at the top of Face 3 
(Text-fig. 1). Beds forming the uppermost meter of 
section on Face 3 contain an abundance of palm pet- 
iolar debris, abraded wood fragments, and small 
rounded fossiliferous chert clasts. The chert clasts, 
mostly 0.5-1.2 cm in diameter, may represent frag- 
ments of peat that were ripped up and carried along 
with the other clasts, or could alternatively represent 
small coprolites (C. B. Hanson, pers. comm.). Leaf 
remains in this upper horizon are usually folded or 
rolled, not flat-lying, and apparently were deposited 
during turbulent, rather than quiet-water conditions. 
The sedimentary environment represented by the 
Nut Beds deposit has been interpreted as that of a lake 
delta deposit (Scott, 1954; C. B. Hanson, pers. comm.). 
This interpretation would account for the large amount 
of jumbled biotic debris, and the general thinning of 
the beds toward the northeast (C. B. Hanson, pers. 
comm.). However, typical lacustrine sediments, with 
fissile shales containing fish remains, as observed at 
other Clarno paleobotanical localities (e.g., West Branch 
Creek, White Cliffs; Manchester, 1990), are not present 
at the Nut Beds. The lower portion of the section, with 


alternating siltstones and sandstones, might represent 
periodic flood deposition, whereas some of the thick, 
poorly sorted upper layers evidently represent a rapidly 
deposited slurry, possibly from the toe of a volcanic 
mudflow. Local hot springs may have been present, 
contributing toward rapid mineralization of biotic de- 
bris. 


AGE 


The value of the Nut Beds flora for biostratigraphic 
and paleobiogeographic correlations is enhanced by an 
accurate determination of its age. In previous paleo- 
botanical assessments, age interpretations for the Clar- 
no Nut Beds have ranged from “older than Upper 
Eocene" (Scott, 1954) to Early Oligocene (Wolfe, 1971, 
1981). Although Wolfe (1981) attributed the Nut Beds 
(“late Clarno") flora to his Kummerian floral stage, of 
inferred Late Eocene/Early Oligocene age, his zonation 
is based on leaf species from a type section in the Puget 
Group of northwestern Washington (Wolfe, 1968). It 
is impossible to determine which of the Clarno fruit 
and seed species might correspond to the leaf species 
that Wolfe used to characterize his stages, and prelim- 
inary investigations of the Nut Beds leaf assemblage 
(Manchester, 1981) indicate that the flora is no more 
similar taxonomically to the Kummerian stage than to 
the older Franklinian and Ravenian stages. In any case, 
to eliminate potential for circularity, the age of the Nut 
Beds flora is best determined by means independent 
of paleobotanical correlation. 

Radiometric dates reported for various parts of the 
Clarno Formation range from about 54 to 34 million 
years (Fiebelkorn et al., 1983), although the younger 
dates are questionable because the basal ignimbrite of 
the overlying John Day Formation has been dated at 
about 37 (Swanson and Robinson, 1968; Fiebelkorn 
etal., 1983; Vance, 1988) to 39.7 (Bestland et al., 1993) 
million years. The Clarno Formation represents a com- 
plex volcanic terrain characterized by abrupt lateral 
and vertical variation complicated by erosion, faulting 
and slumping. Therefore, it is difficult to trace indi- 
vidual units beyond local areas, and, as a result, the 
stratigraphic position of the Nut Beds in relation to 
other datable units of the Clarno Formation is not 
immediately obvious. Fortunately, the Nut Beds de- 
posit itself contains a vertebrate fauna that can be cor- 
related with North American land mammal stages, as 
well as volcanic clasts suitable for radiometric dating. 

Stirton (1944) described a tooth of Hyrachyus from 
the Nut Beds, and on this basis clearly established an 
Eocene age for the locality. More complete cranial 
specimens of Hyrachyus, along with remains of five 
other mammalian vertebrates, including Orohippus, 
Patriofelis and Telmatherium, have been investigated 


| 
| 
| 
| 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 13 


by Hanson (1973) and clearly represent a Bridgerian 
fauna (Hanson, pers. comm., 1988). This fauna occurs 
within the Nut Beds along with the fruits and seeds, 
and should not be confused (cf., Wolfe, 1981, p. 44) 
with the younger Clarno fauna from Hancock Quarry, 
1 km northeast of the Nut Beds, that is of Chadronian 
or Duchesnian age (Mellet, 1969; Hanson, 1973, pers. 
comm., 1988). By correlation with radiometrically 
dated faunas of the Rocky Mountain region, the Bridg- 
erian vertebrate fauna of the Nut Beds indicates a Mid- 
dle Eocene age of about 45 m.y. 

Although a potassium-argon date of 34 million years 
was published for the Nut Beds (Evernden et al., 1964), 
such a young age (Late Eocene or Early Oligocene) is 
very doubtful because the basal ignimbrite of the over- 
lying John Day Formation (situated about 30 meters 
Stratigraphically above the Nut Beds) yields radio- 
metric dates of about 37 (potassium-argon, Swanson 
and Robinson, 1968; fission track, Vance, 1988) to 
39.7 (argon-argon, Bestland et al., 1993) million years. 
More recently, Vance (1988) obtained fission track dates 
of 43.6 and 43.7 (+ 10%) m.y. based upon zircon 
extracted from Nut Beds sediment. In 1987, C. B. Han- 
son, B. D. Turrin and I collected a sample of water- 
reworked tuff with intact pumice clasts from a plant 
and bone-bearing stratum at the middle stratigraphic 
level of Face 1 in the Nut Beds. Ten plagioclase crystals 
isolated from this sample were analyzed by the single 
crystal laser fusion argon 40/39 technique and yielded 
dates ranging from 36.38 + 1.31 to 46.8 + 3.36 my. 
(B. D. Turrin, pers. comm., 1988). Statistically, these 
dates give an arithmetic mean of 42.98 + 3.48 witha 
Standard error of the mean, + 1.10, and a weighted 
average of 43.76 + 0.29 m.y., strikingly close to the 
fission track results of Vance (1988). Based upon the 
recently obtained radiometric ages and the vertebrate 
Correlations, the age of the Nut Beds flora may be 
considered to be Middle Eocene, within the range of 
43 to 45 million years old. 


OTHER CLARNO FLORAS 


Although this report is intended to be as compre- 
hensive as possible for the fruit and seed assemblage 
of the Nut Beds locality, it should not be regarded as 
an exhaustive treatment of the Clarno flora. The Clar- 
по Formation occurs over an area of about 4000 km2 
(Walker and Robinson, 1990) and includes many fossil 
Plant localities (Text-fig. 2). The location and general 
Character of some of these localities are reviewed by 
Hergert (1961) and Manchester (1990). Geographic co- 
Ordinates for localities from which large collections 
have been made have been published previously (Man- 
Chester, 1986, p. 224; 1991, p. 717). Some of these 
Sites, particularly the lacustrine deposits in the West 


Branch Creek and Cherry Creek drainages, include 
many species of leaves and compressed fruits that are 
not known from the Nut Beds. I have made reference 
to these other localities where appropriate in discussing 
the distribution of individual species. 

Another occurrence of fossil fruits and seeds in the 
Clarno Formation is the Late Eocene (Duchesnean) 
Hancock Quarry vertebrate locality, situated 1 km 
northeast of the Nut Beds. This locality (UF 70) has 
provided abundant fruit and seed remains preserved 
as partially compressed carbonaceous casts and molds 
in claystone (McKee, 1970). Although anatomical de- 
tails are not preserved, external morphology enables 
identification through comparison with the better pre- 
served Nut Beds specimens. More than 500 specimens 
were collected from this site (collections at OMSI and 
UF) and is possible to distinguish about 30 taxa. Sev- 
eral genera are shared with the Nut Beds, including 
Ampelocissus, Diploclisia, Juglans, Odontocaryoidea, 
Palaeophytocrene, and Vitis. In addition to the genera 
recognized by McKee, I have observed specimens of 
Alangium, Eohypserpa, Iodes, Iodicarpa, Mastixioi- 
diocarpum and Pentoperculum in the Hancock Quarry 
assemblage. Thus, it is clear that some of the Nut Beds 
genera persisted in the same region at least until the 
Late Eocene. There are, however, marked floristic dif- 
ferences between both of these assemblages and the 
Lower Oligocene Bridge Creek flora of the overlying 
John Day Formation (Chaney, 1927; Manchester and 
Meyer, 1987; Manchester, 1990). 

The Brummers Spring locality (UF 254), also in the 
Clarno Formation, is located about 90 km south of the 
Nut Beds locality and 20 km east of Post, Oregon 
(Text-fig. 2). This locality has produced a small assem- 
blage of silicified fruits and seeds including Juglans, 
Magnolia, Mastixicarpum, Quercus, and Sabal. 


MODES OF PRESERVATION 


Fossil fruits and seeds in the Nut Beds are preserved 
in various modes, ranging from compressions and im- 
pressions to molds and casts, to permineralizations. 
These different. types of preservation, as applied to 
fossil plant tissues, are reviewed by Scott and Collinson 
(1983, p. 118). Compressed fruits and seeds, preserved 
along with leaf impressions near the base of the section 
(Manchester, 1981), usually lack internal structural de- 
tails, but are useful in providing characters of wing 
morphology and venation. Three-dimensionally pre- 
served specimens, including casts, molds and permi- 
neralizations, occur throughout the deposit. 

Casts and molds lacking internal anatomy are com- 
mon (Scott, 1954), but sometimes the internal tissues 
are preserved through permineralization by chalced- 
ony and/or calcite. A single specimen may include more 


14 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


than one mode of preservation; for example, a cast of 
the locule may occur within the carbonized or per- 
mineralized remains of an endocarp. Permineralized 
seed remains are sometimes encountered within the 
locule cast. 

Silicification appears to have been primary, and cal- 
cification secondary, as in the case of walnuts that are 
sometimes preserved with the locule filled by chalced- 
ony and the nutshell infiltrated by calcite. The quality 
of preservation in permineralized specimens is vari- 
able, depending on porosity and durability of the tis- 
sues involved. Fleshy or pulpy outer fruit layers are 
rarely if ever preserved. Woody layers tend to be well- 
preserved, sometimes with excellent cellular detail. 
However, thin, hard, evidently impermeable layers 
typically are represented only as an airspace between 
the chalcedony internal mold and the siltstone outer 
mold of the structure, as in the case of seeds of An- 
nonaspermum, Fortunearites, and Bumelia. 


MATERIALS AND METHODS 


Approximately 20,000 specimens were examined for 
this study. The majority of specimens were collected 
between 1942 and 1983 through the extraordinary ef- 
forts of Thomas J. Bones. Additional specimens were 
collected in the 1940’s and 50’s by A. W. Hancock, R. 
A. Scott, and, during the summers of 1974-1989, by 
S. R. Manchester with the help of many high school 
students (acknowledged p. 7) in conjunction with pa- 
leobotanical programs at Hancock Field Station, a nat- 
ural science facility operated by the Oregon Museum 
of Science and Industry 1.5 km southeast of the Nut 
Beds. 

The fossil disseminules range from less than 1 mm 
to 85 mm in length. Because the sediment is silica- 
cemented, standard washing and sieving techniques 
(Tiffney, 1990) are not effective in recovering speci- 
mens. Instead, the fossils are obtained through a labor- 
intensive process of prying out blocks of sediment and 
breaking the rock with hammers. Fruits and seeds are 
exposed in the matrix when intercepted by the resulting 
fracture planes. The percussion frequently causes the 
fruit or seed to pop out of the matrix. Sometimes outer 
layers of the fossil remain attached to the matrix mold. 
It is thus important to retain the associated mold for 
study as well as the more attractive cast. Thomas Bones 
collected large numbers of small seeds by crushing the 
matrix and sorting with the aid of a magnifying lens. 

Because the type and quality of preservation varies 
greatly, even among specimens of the same species, it 
was necessary to sort the specimens under a dissecting 
microscope to recover the most informative samples 
for describing each species. Silica casts and molds re- 


placing various tissues of the fruit or seed were useful 
for morphology and dimensions. Permineralized spec- 
imens were most valuable in providing the best details 
of internal morphology and anatomy. Permineraliza- 
tions were not available for all species, but in many 
instances a careful examination of numerous speci- 
mens resulted in the recovery of at least a few per- 
mineralized examples that could be fractured or sawn 
to reveal internal morphology and anatomy. Com- 
pression and impression specimens were also found to 
be useful because they enable direct comparison with 
specimens from other Clarno localities where permi- 
neralizations are not available, and because they pro- 
vide details of wing morphology and venation for sam- 
aras that usually are not evident in the permineralized 
specimens. 

Morphology and anatomy was recorded photograph- 
ically with Kodak Technical Pan 2415 film processed 
for medium contrast. Lighting for reflected light pho- 
tography was provided from the upper left and lower 
right side of each specimen with a pair of small, diffuse 
tungsten lamps that could be oriented at various angles 
to provide optimal illumination of surface detail. Spec- 
imens of medium to large size (>8 mm diam.) were 
photographed with a Nikon FE camera with 100 mm 
Micronikkor lens. Macrography of smaller specimens 
(0.5-8 mm) was conducted with the FE camera mount- 
ed on a Nikon SMZ10 zoom stereo scope. Stereo-pair 
photographs were prepared of specimens which were 
subsequently fractured or serially sectioned. For ste- 
reo-pairs, successive pictures were taken with the spec- 
imen rotated 6 degrees on a teetering stand, or by using 
the stereo-photo option on the Nikon SMZ10 macro- 
scope. The translucency and differential coloration of 
silicified specimens initially presented problems, ob- 
scuring surface details in photography. These problems 
were overcome by coating the specimens with palla- 
dium using a sputter coater prior to light micrography- 
The palladium was subsequently removed by soaking 
for 30 minutes in a saturated solution of sodium cy- 
anide in water (Sela and Boyd, 1977). Small seeds, and 
minute surface details of larger specimens, were stud- 
ied by standard techniques of scanning electron mi- 
croscopy (SEM), using the Cambridge Stereoscan 250 
and Hitachi S-450 SEM models. 

Internal morphology and anatomy were revealed in 
some instances by studying fractured specimens with 
light and SEM microscopy and in other cases by serial 
sectioning. Specimens 1.5 mm to 25 mm in diameter 
were sectioned with a Microslice II annular diamond 
saw, with a very thin blade (0.05 mm thick) to mini- 
mize wastage (kerf loss about 0.15 mm) in the cutting 
process. Resulting wafers were photographed unetched 
in xylene, or dry after etching with hydrofluoric acid, 


| 
| 


| 
| 
| 


—————————— 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 15 


by reflected light microscopy using a Wild M400 pho- 
tomacroscope. For finer anatomical details the wafers 
were then used to make peels and/or ground thin sec- 
tions for transmitted light microscopy. Peels were pre- 
pared by the cellulose acetate peel method (Basinger 
and Rothwell, 1977; Basinger, 1981) after etching for 
two minutes in 48% HF. The peels were cleared in 
clove oil, mounted in Canada Balsam with slide and 
coverglass. Ground sections were prepared by mount- 
ing Ше wafers on microscope slides with Elmers Epoxy, 
and grinding to approximately 30 um thickness on a 
Buehler thin sectioning machine. Anatomical detail 
from thin sections was photographed with the Nikon 
FE camera mounted on a Nikon Labophot compound 
microscope using standard transmitted and/or reflect- 
ed light from an obliquely positioned fiber optic source. 
For one taxon (Hydrangea), small organically pre- 
served seeds were liberated by dissolving the silicified 
fruit in hydrofluoric acid, then washed and mounted 
on microscope slides for transmitted light microscopy. 

External measurements were obtained with brass 
calipers graduated to 0.1 mm. Dimensions were cal- 
culated on the basis of all available specimens, or at 
least 25 specimens, of each species. Obviously con- 
torted or broken specimens were not included in the 
measurements. Unless otherwise indicated, the longest 
axis of the fruit or seed was treated as height or length. 
Width and thickness are the wider and narrower di- 
mensions, respectively, measured at right angles to the 
length. 

Methods of identifying fossil fruits and seeds are 
reviewed in detail by Tiffney (1990). Discussions of 
€xtant and fossil seeds by Reid and Chandler (1933), 
Chandler (1961b, 1964, 1978), Collinson (1983), Friis 
(1985), Kirchheimer (1957), and Mai (1976) provided 
à useful guide for preliminary investigations of the 
Clarno taxa. Comparisons with extant seeds were car- 
ried out by examining representatives of most extant 
angiosperm families in herbaria and seed collections. 


RESULTS AND INTERPRETATION 


SYSTEMATIC DIVERSITY OF THE 
NUT BEDS FLORA 


One hundred and forty five genera and 173 species 
Of fruits and seeds have been recognized among the 
Nut Beds collections (Table 1) and are described and 
illustrated in this monograph. In addition, among the 
less well-preserved specimens in the collections, there 
mày be as many as twenty more genera that are ex- 
Cluded from the following discussions. Four gymno- 
Sperm genera, representing Pinaceae and Taxaceae, are 
recognized, but the majority of taxa are angiosperms. 


From the working total of 145 genera and 173 species, 
only 75 genera and 102 species have been identified 
as belonging to modern families. The remaining taxa, 
constituting approximately half of the assemblage, are 
of uncertain affinity with respect to extant families 
(Text-fig. 5A). This group probably includes both ex- 
tant and extinct genera, but the affinities are still un- 
determined at the time of this writing. Nevertheless, 
these species are described and illustrated in order to 
document, as completely as possible, the full diversity 
of the flora and to invite further study by other re- 
searchers. 

The numbers presented above, based only on the 
fruits and seeds, understate the full diversity of the Nut 
Beds flora because many additional taxa are repre- 
sented by other plant organs. For example, Equisetum 
clarnoi (J. T. Brown, 1975; Retallack, 1981) is known 
from silicified stems and compressed strobili (Pl. 1, fig. 
1); four types offerns are known from leaves and stems, 
and a cycad with foliage resembling Dioon is present 
(Manchester, 1981). Families confirmed on the basis 
of wood that have not been recognized among the fruits 
and seeds include Ginkgoaceae (Ginkgo; Scott et al., 
1962), Trochodendraceae (Scott and Wheeler, 1982), 
and Sterculiaceae (Manchester, 1979, 1980). Although 
the extent of overlap is unknown, it is probable that 
the fossil leaves (Manchester, 1981) and woods (Scott 
and Wheeler, 1982) collected from the Nut Beds rep- 
resent some of the same species as those known from 
seeds or fruits treated here. However, because none of 
the organs have been found in direct attachment, as- 
sessments of conspecificity remain speculative. 

It is not possible to give a precise percentage of the 
Nut Beds genera that are extinct vs. extant because a 
large proportion of the genera are still of uncertain 
systematic position relative to modern families and 
genera. Nevertheless, all genera recognized here may 
be divided into four groups indicating current knowl- 
edge of their status relative to living taxa (Text-fig. 5): 

1) Extant genera. Those that possess features that 
are diagnostic of a living genus; these species are given 
the appropriate modern generic name, for example, 
Cornus, Magnolia, Meliosma, Quercus. 

2) Extinct genera. Those that have features diag- 
nostic of a particular modern family, but with addi- 
tional characters that set them apart from living genera 
of that family, for example, Cruciptera, Langtonia, 
Pentoperculum. 

3) Form genera. Those that have not been traced to 
a particular modern family (i.e., incertae sedis), for 
example, Carpolithus, Joejonesia. Form genera prob- 
ably include extant genera that I have failed to identify, 
as well as extinct genera and/or families. These genera 


16 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Table 1.—Comprehensive list of fruit and seed taxa from the Clarno Nut Beds assemblage indicating number of specimens observed. 


taxon quantity taxon quantity 
Pinaceae Palaeophytocrene pseudopersica Scott 80 
Pinus sp. 1 Pyrenacantha occidentalis sp. nov. 8 
Taxaceae Juglandaceae 
Taxus masonii sp. nov. 20 Juglandeae 
Torreya clarnensis sp. nov. qs Cruciptera simsonii (Brown) Manchester 12 
Diploporus torreyoides gen. et sp. поу. 5 Juglans clarnensis Scott 200+ 
Actinidiaceae Engelhardieae 
Actinidia oregonensis sp. nov. 7 cf. Palaeocarya clarnensis Manchester 48+ 
Alangiaceae Platycaryeae 
Alangium eydei sp. nov. 33 Paleoplatycarya? hickeyi sp. nov. 5 
Alangium rotundicarpum sp. nov. 1 Ташасоле 
Anacardiaceae Laurocalyx wheelerae sp. nov. 12 
Pentoperculum minimus (Reid et Chandler) Laurocarpum hancockii sp. nov. 1 
gen. et comb. nov. 62 Laurocarpum nutbedensis sp. nov. 1 
Rhus rooseae sp. nov. 15 Laurocarpum raisinoides sp. nov. 1 
Аппопасеае Lindera clarnensis sp. nov. 8 
Anonaspermum cf. pulchrum Reid et Chandler 16 Leguminosae 
Anonaspermum bonesii sp. nov. 1 Leguminocarpon sp. 1 
Anonaspermum rotundum sp. nov. 1 Lythraceae 
Araliaceae Decodon sp. 7 
Paleopanax oregonensis gen. et 5р. поу. 3 Magnoliaceae 
Betulaceae Magnolia muldoonae sp. nov. 100+ 
Kardiasperma рагуит gen. et sp. поу. 21 Magnolia paroblonga sp. nov. 17 
Coryloides hancockii gen. et sp. поу. 34 Magnolia tiffneyi sp. nov. 10 
Burseraceae Е 
: Menispermaceae 
Bursericarpum oregonense Sp. поу. 40 ; 
Coscineae 
Bursericarpum sp. 6 : џ 
Anamirta leiocarpa sp. поу. 16 
Cornaceae р 
; Menispermeae 
Cornus clarnensis sp. nov. 5 + de д } a 
Langtonia bisulcata Reid et Chandler 31 Diploclisia aurif С (Hollick) comb. nov. 8 
Mastixia sp. 4 Eohypserpa scottii sp. nov. 2 
Mastixioidiocarpum oregonense Scott 22 Davisicarpum limacioides зразок 1 
Mastixicarpum occidentale sp. nov. 2 Palaeosinomenium venablesii Chandler 70 
Nyssa scottii sp. nov. 6 Tinosporeae 
Nyssa spatulata (Scott) comb. nov. 29 Atriaecarpum clarnense sp. nov. 4 
Nyssa sp. 5 Calycocarpum crassicrustae sp. nov. 40 
Fagaceae Chandlera lacunosa Scott 40 
Castanopsis crepetii sp. nov. 6 Curvitinospora formanii gen. et sp. nov. 1 
Quercus paleocarpa Manchester 3 Odontocaryoidea nodulosa Scott 80+ 
: Tinospora elongata sp. nov. 7 
Flacourtiaceae a 
Е Е Tinospora hardmanae sp. nov. 4 
Saxifragispermum tetragonalis sp. поу. 65 Tinomiscoidea occidentalis sp. nov. 1 
Hamamelidaceae Thanikaimonia geniculata gen. et sp. nov. 1 
Fortunearites endressii gen. et sp. поу. 6 Musaceae 
Hydrangeaceae Ensete oregonense Manchester et Kress 70 
Hydrangea knowltonii gen. et sp. nov. 45+ Palmae 
Icacinaceae Sabal bracknellensis (Chandler) Mai 6 
Iodeae Sabal jenkinsii (Reid et Chandler) comb. nov. 30 
Todes multireticulata Reid et Chandler 14 Platanaceae 
Iodes chandlerae sp. nov. 13 Macginicarpa glabra Manchester 200+ 
Iodicarpa ampla gen. et sp. nov. ди Platanus hirticarpa sp. nov. 1 
Iodicarpa lenticularis sp. nov. 16+ Tanyoplatanus cranei gen. et sp. поу. 3 
Comicilabium atkinsii gen. et sp. nov. 8 Ёозассаё 
Phytocreneae Prunus weinsteinii sp. nov. 1 
Palaeophytocrene hancockii Scott 20 Prunus olsonii sp. nov. 10 


| 
| 
| 
| 
| 
| 


Hexacarpellites hallii gen. et sp. nov. 
Joejonesia globosa gen. et sp. nov. 


Unknown families: 70 genera, 71 species 
Total: 145 genera, 173 species 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER d 
Table 1.— Continued. 
taxon quantity taxon quantity 
Rubiaceae Lignicarpus crassimuri gen. et sp. nov. 4 
Emmenopterys dilcheri sp. nov. 7 Ligniglobus sinuosifibrae gen. et sp. nov. 5 
Sabiaceae Lunaticarpa curvistriata gen. et sp. nov. 80 
Meliosma beusekomii sp. nov. 110 Microphallus perplexus gen. et sp. nov. 23 
Meliosma bonesii sp. nov. 22 Nephrosemen reticulatus gen. et sp. поу. Zi 
Meliosma elongicarpa sp. nov. 2 Omsicarpium striatum gen. et sp. пом. 1 
Meliosma cf. jenkinsii Reid et Chandler 8 Pasternackia pusilla gen. et sp. nov. 220 
Meliosma leptocarpa sp. nov. 50+ Pileosperma minutum gen. et sp. nov. 10 
Sabia americana sp. nov. 22 Pileosperma ovatum sp. nov. За 
Sapindaceae Pistachioides striata gen. et sp. nov. 2 
Devincer ин aren: 28 Pollostosperma dictyum gen. et sp. nov. 260 
Palaeoallophylus globosa sp. nov. 36 — аи age acti: "D я 
Palaeoallophylus gordonii sp. nov. 20 TR ким ас први ^ 
MEN Pteronepelys wehrii gen. et sp. nov. 2 
Pulvinisperma minutum gen. et sp. nov. 170 
Bumelia? globosa sp. nov. 40+ Pyrisemen attenuatum gen. et sp. nov. 32 
Bumelia? subangularis sp. nov. B Quintacava velosida gen. et sp. nov. 1 
Schisandraceae Sambucuspermites rugulosus gen. et sp. nov. 9 
Schisandra oregonensis sp. nov. 27 Scabraecarpium clarnense gen. et sp. nov. 9 
Staphyleaceae Scalaritheca biseriata gen. et sp. nov. 10 
Tapiscia occidentalis Manchester 78 Scaphicarpium radiatum ant span. 13 
EN Sphaerosperma riesii gen. et sp. nov. 2 
Sphenosperma baccatum gen. et sp. nov. 1 
Symplocos nooteboomii sp. nov. d Stockeycarpa globosa gen. et sp. nov. 1 
Theaceae Striatisperma coronapunctatum gen. et sp. nov. 69 
Cleyera grotei sp. nov. 32 Tenuisperma ellipticum gen. et sp. nov. 2 
RA Tiffneycarpa scleroidea gen. et sp. nov. 1 
ми Trigonostela oregonensis gen. et sp. nov. 9 
Celtidoideae Tripartisemen bonesii gen. et sp. nov. 270+ 
Aphananthe тай sp. nov. 12 Triplascapha collinsonae gen. et sp. nov. 4 
Celtis burnhamae sp. nov. 100+ Triplexivalva rugata gen. et sp. nov. 12 
Celtis sp. 160+ Trisepticarpium minutum gen. et sp. nov. 1 
Trema nucilecta sp. nov. 46 Truncatisemen sapotoides gen. et sp. nov. 3 
Ulmoideae Ulosperma hardingae gen. et sp. nov. 4 
Cedrelospermum lineatum (Lesq.) Manchester 11 Wheelera lignicrustae gen. et sp. nov. 3 
Vitaceae Carpolithus bellispermus Chandler 100 
Е у Carpolithus sp. 1 > 
Ampelocissus auriforma sp. nov. 90 Carpolithus sp. 2 2 
Ampelocissus scottii sp. nov. 10 Carpolithus sp. 3 3 
Ampelopsis rooseae sp. nov. 260 Carpolithus sp. 4 1 
nn... angustisulcata Scott 10 Carpolithus sp. 5 Э 
en clarnensis sp. nov. 32+ Carpolithus sp. 6 1 
itis magnisperma Chandler 9 Carpolithus sp. 7 3 
Vitis tiffneyi sp. nov. 50+ Carpolithus sp. 8 1 
Incertae Sedae Carpolithus sp. 9 10 
Ankistrosperma spitzerae gen. et sp. nov. 48 Carpolithus sp. 10 4 
Anonymocarpa ovoidea gen. et sp. nov. 1 Carpolithus sp. 11 1 
Ascosphaera eocenis gen. et sp. поу. 20 Carpolithus sp. 12 1 
Axinosperma agnostum gen. et sp. nov. 22 Carpolithus sp. 13 1 
Bonesia spatulata gen. et sp. nov. 15 Carpolithus sp. 15 1 
Comminicarpa friisae gen. et sp. nov. 5+ Carpolithus sp. 16 15 
Cuneisemen truncatum gen. et sp. nov. 9 Carpolithus sp. 17 1 
Dentisemen parvum gen. et sp. nov. 3 Carpolithus sp. 18 2 
Durocarpus cordatus gen. et sp. nov. 1 Carpolithus sp. 19 1 
Ferrignocarpus bivalvis gen. et sp. поу. 24 Carpolithus sp. 20 1 
Fimbrialata wingii gen. et sp. nov. 10 Carpolithus sp. 21 1 
Fragarites ramificans gen. et sp. поу. 7 Five-part flower 2 
Globulicarpium levigatum gen. et sp. nov. 20 Extant families: 75 genera, 102 species 


18 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


include some interesting puzzles for future systematic 
research. 

4) Stereotype genera. Those that have features di- 
agnostic ofa particular modern family, with a suite of 
characters found in more than one living genus of that 
family. Species assigned to stereotype genera cannot 
be resolved to a single extant or extinct genus; for 
example, Anonaspermum, Laurocarpum, Palaeosino- 
menium. 

One must be cautious when interpreting fossil ge- 
neric names. Form genera should not be blanketly as- 
sumed to represent extinct genera, because it is just as 
likely that they may represent undetermined extant 
genera. Likewise, in the characters available for study, 
stereotype genera are indistinguishable from two or 
more extant genera, and accordingly should not be 
assumed to represent extinct genera. 

The Nut Beds biota also includes insect remains. 
Two types of beetles are known from silicified adults 
(5. В. Manchester, pers. obs.) and one of Ше taxa pre- 
liminarily thought to be silicified seeds has been de- 
termined to represent eggs of the Phasmidae (Sellick, 
in press). The possibility exists that some of the form 
genera described in this monograph represent insect 
eggs or egg cases rather than plant remains. Rather 
than exclude taxa that could possibly be construed as 
insect eggs, I believe it is useful to document all of the 
structures because of the possible importance for bio- 
stratigraphy and/or future insect work. As this goes to 
press, I consider the following taxa worthy of inves- 
tigation as possible insect remains: Carpolithus bellis- 
permus, Scalaritheca and Pileospermum. These taxa, 
nevertheless, have been included in the fruit/seed counts 
and, if ultimately determined to be of insect origin, 
will slightly alter some of the percentages presented in 
this section. 

Thirty-five extant families have been recognized 
among the Nut Beds fruits and seeds. Among the 75 
genera of known familial affinities, 38, or 5096, have 
been attributed to genera that are still living today 
(Text-fig. 5B). These extant genera, indicated in Tables 
1 and 5, are significant in showing little or no evolution 
in fruit morphology over a period of at least 43 million 
years. The number of extant genera recognized may be 
expected to increase with additional work as the аћп- 
ities of form genera come to light. 

At least 26 genera (3596 of those for which familial 
affinities have been determined) are extinct: Atriae- 
carpum, Cedrelospermum, Chandlera, Comicilabium, 
Kardiasperma, Coryloides, Cruciptera, Curvitino- 
spora, Davisicarpum, Deviacer, Diploporus, Eohyp- 
serpa, Fortunearites, Iodicarpa, Langtonia, Mac- 
ginicarpa, Mastixicarpum, Mastixioidiocarpum, 
Odontocaryoidea, Palaeophytocrene, Palaeosinomen- 
ium, Pentoperculum, Saxifragispermum, Tanyopla- 


tanus, Thanikaimonia, Tinomiscoidea. This figure, too, 
will probably change with additional work. Sometimes 
fossils initially considered to be extinct are subsequent- 
ly found to be living as neobotanical exploration and 
comparative work continues, as in the classic case of 
Metasequoia, and as more recently discovered in Di- 
plopanax (Eyde and Xiang, 1991) and Craigia (Kva- 
éek, Büzek and Manchester, 1991). Morphological dif- 
ferences between related fossil and extant taxa that may 
be viewed as warranting generic distinction by one 
investigator may be viewed as simply reflecting greater 
variability within a single genus by another. For ex- 
ample, the species formerly treated as Palaeonyssa by 
Reid and Chandler (1933) and Scott (1954) are herein 
regarded as Nyssa. 

Relative to other paleobotanical localities of the 
Clarno Formation, the Nut Beds fruit and seed assem- 
blage is anomalously high in generic and specific di- 
versity. In surveying the large collections from the la- 
custrine shales at West Branch Creek (UF loc. 229c, 
230, UCMP loc. 3904, USGS loc. 8637), I have ob- 
served only 48 distinct types of compressed fruits and 
seeds and 53 types of leaves. 

The leaf flora of the Nut Beds includes about 65 
genera—less than half as many as those known from 
fruits and seeds. Although the leaf collection acquired 
from the Nut Beds between 1974 and 1981 1s the result 
of more than 2000 person-hours, the recovery process 
for leaves in the heavily fractured siltstones (Man- 
chester, 1981) is very time-consuming and the total 
number of leaf specimens is less than 1000. The fruit 
and seed collections amassed since 1943 include more 
than 20,000 specimens. Thus, the discrepancies be- 
tween foliage and diaspore diversity may be in part 
due to differences in sample size. 

It's unlikely that all of the Nut Beds species grew 
together in the same plant community; rather, it is 
probable that the assemblage includes elements from 
various parts of a large watershed. Judging from the 
sedimentary environment of the Nut Beds and the 
varying amounts of predepositional erosion evident in 
the fossil remains, the assemblage probably includes 
plants from different sources: some local, others carried 
from various distances upstream. Significant transport 
of fruits and seeds may also have been effected by birds 
that consumed fruits from various parts of the forest. 
When perched on branches close to or overhanging 
sites of sediment accumulation, birds have the poten- 
tial to introduce endocarps and seeds from more re- 
mote parts of the forest that would otherwise be un- 
derrepresented in the fossil assemblage. Such biotic 
transport, selective for bird-edible fruits, might explain 
the greater diversity of fruits and seeds than vegetative 
organs in the Nut Beds. Frugivorous mammals prob- 
ably also have played a role in fruit dispersal in Eocene 


| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 19 


Unknown 


48% 


Modern Genera, 
Stereotype Genera, | 
Extinct Genera 


Form Genera 


48% 


Families 


52% 


15% 


Stereotype genera (11) 


50% 


Ф 
г. 
+ 
oe 7 
vr — 
» 
* 
2 


7 35% 


7 


PAPIERE 
Ci d Pu 


2 


Text-figure 5.— Taxonomic status of Nut Beds fruit and seed genera relative to extant families and genera. A, Proportion of genera identified 
to modern families vs. those of unknown familial affinities. B, Genera identified to modern families, with proportions of extinct, modern and 
Stereotype genera. C, Summary, emphasizing that only У, of genera have been determined to be extant. 


forests (Collinson and Hooker, 1992). Indeed, a large 
Proportion of the seeds and endocarps identified ap- 
Dear to have been from drupes or berries that would 
have been well-suited for bird and/or mammal dis- 
Persal. The apparent absence of vitaceous leaf remains 
In the Clarno collections, contrasted with the abun- 


dance and diversity of grape seeds, supports this con- 
clusion. 


GROWTH HABIT AND DISPERSAL ECOLOGY 


| The growth habit of a fossil species, i.e., whether it 
15 a tree, shrub, Папа or herb, may be inferred from 
that of its living relatives. Of course, this means of 
Inference is not applicable to the fruit and seed taxa 
whose modern affinities are undetermined; therefore, 
fewer than half of the Nut Beds fruit and seed genera 
Сап be analyzed according to this approach. Never- 
theless, this methodology gives some insight into the 


stature of Nut Beds vegetation (Table 2). Among the 
identified taxa, trees and lianas are particularly well 
represented, while herbs and aquatic plants are poorly 
represented. 

Many of the fruit and seed genera are deduced to 
represent trees. Arborescent taxa include Aphananthe, 
Castanopsis, Juglandaceae (3 genera), Lauraceae, Mac- 
ginicarpa, Magnolia, Mastixia, Meliosma, Nyssa, 
Platanus, Quercus, Sabal, Tapiscia. Trees of lower 
stature and shrubs probably included Annonaceae, 
Cornus, cycad (known from foliage only), Fortunear- 
ites, Taxus, and Torreya. Arborescent taxa can also be 
inferred independently of fruit and seed data using 
fossil wood fragments from the Nut Beds. Based on 
curvature of growth rings, at least 40 different genera 
were derived from stems larger than 15 cm in diameter 
and were probably from trees. Some of these woods 
have been identified, including: Ginkgo (Scott et al., 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Table 2.—Present habit and vegetational distribution of Nut Beds genera. 


deciduous/ 

genus habit evergreen TRF PTRF NBLE MM MBLD MNH MC 
Actinidia с а + + + ar ar 
Alangium tsc de s + + + Es 
Ampelocissus (e d de 
Ampelopsis d dE ES 
Anamirta © e + + 
Aphananthe $ de at + ES ar 
Bumelia ts d + + 
Calycocarpum с а + 
Castanopsis ts e d dE 3E ar 4E 15 + 
Celtis t de + + + F + 
Cleyera s e + 
Cornus ts d Qu + 3r di 
Decodon S d ar ar 
Diploclisia с е 3pm + 
Emmenopterys t d T 
Ensete h e ae Es 
Hydrangea tsc de T + + + + + 
Iodes с e Е НЕ 
Juglans t + + + + + 
Lindera ts d ES + AB qe 
Magnolia t de + НЕ + + + dr 
Mastixia t e e 
Meliosma t de eJ 4 ar ar Н + 
Parthenocissus с а ar 
Pinus t e 7 ae Ae dE qe 4E Ar 
Platanus t d + + + + 7 
Prunus ts de К dp EN 3E ar 
Pyrenacantha С ё ap dc ar 
Quercus + де + ap 4E зБ dis dE jr 
Rhus tsc de +1 s. E 4r 
Sabal + е ar + + 
Sabia с e ar 
Schisandra © de dE ES + 
Symplocos ts de + + + + + + 
Тарїзсїа ts d dE 
Taxus t @ Ar a 
Tinospora g © 46 + 
Torreya t e Ae E 
Trema ts e ap dE 4p 
Vitis с а T de a Ar 


' Including submontane rainforest. 
2 Including montane forest. 


c = climber, s = shrub, t = tree, h = herb, d = deciduous, e = evergreen; TRF = Tropical Rain forest; PTRF = Paratropical Rain forest; 
NBLE = Notophyllous Broadleaved Evergreen; MM = Mixed Mesophytic; MBLD = Mixed Broadleaved and Deciduous; MNH = Mixed 
Northern Hardwood; MC = Mixed Coniferous. Vegetation types following Wolfe (1979). 


1962), Tapirira (Manchester, 1977), Sterculiaceae 
(Manchester, 1979, 1980), Alangium, Betula, Faga- 
ceoxylon, Magnolia, Plataninium, Quercinium, Ul- 
minium, (Scott and Wheeler, 1982), Engelhardioxylon 
(Manchester, 1983), and Clarnoxylon (Manchester and 
Wheeler, 1993). 

The diversity of climbers in the flora, at least 22 
genera and 30 species, (43% ofthe 69 species for which 
growth form may be inferred) is particularly striking. 
This estimate is based upon extant genera of climbing 
habit (Table 2), plus extinct genera belonging to fam- 
ilies or subfamilies that are virtually all climbers today. 


Menispermaceae, a family comprised almost exclu- 
sively of tropical lianas, is the most diverse family in 
the Nut Beds fruit and seed assemblage, with 13 genera 
and 14 species. In addition, seven species of Vitaceae, 
at least five species of Icacinaceae (those belonging 10 
Iodes, Iodicarpa, Palaeophytocrene, Pyrenacantha), and 
the species of Actinidia, Sabia, and Schisandra prob- 
ably represent lianas or climbers (Table 2). By contrast 
with the Nut Beds figure of 43%, the highest known 
percentages of liana species per flora in extant vege- 
tation are only about 24% (Gentry, 1992). 

Lianas such as those of extant Menispermaceae are 


| 
| 
| 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 21 


best developed where the tree canopy is interrupted, 
for example along stream banks and forest edges (For- 
man, 1986). Preferential growth at stream side, where 
seeds may be dropped directly into the sedimentary 
environment, may have caused over-representation of 
lianas relative to trees, shrubs and herbs in the fossil 
assemblage, but I doubt that this bias fully explains 
the high apparent diversity oflianas. The same families 
of lianas are also well represented in the Eocene Lon- 
don Clay flora of England (Chandler, 1961b; Collinson, 
1983). The diversity oflianas along with the trees men- 
tioned above may be taken as an indication that a 
multistratal, closed canopy forest had developed (Up- 
church and Wolfe, 1987). In addition, Crane (1987) 
has suggested that lianas and opportunistic shrubs may 
have played a more dominant role in colonizing bare 
ground in the Early Tertiary than today because the 
major radiation of extant angiosperm herbs occurred 
later in the Tertiary. 

Very few ofthe known Nut Beds taxa represent herbs. 
Among Ше fruit and seed genera, Ensete is the only 
known herbaceous element. Stem and leaf records at- 
test to the presence of Equisetum clarnoi (Pl. 1, fig. 1) 
and a few ferns. The ferns are represented by silicified 
petioles of Dennstaedtiopsis and Acrostichum and by 
foliage of Thelypteris iddingsii (Knowlton) MacGinitie, 
and “Asplenium” hurleyensis Berry (Manchester, 1976). 
Lygodium kaulfussii Heer, which is common at many 
Clarno localities (Manchester and Zavada, 1987), has 
not been recovered from the Nut Beds. Floating aquat- 
ic plants such as Nymphaeaceae, Ceratophyllum, Tra- 
ра and Salvinia are absent. Decodon, with a single 
€xtant species that grows as a sprawling shrub at the 
edge of swamps, is the only genus of the Nut Beds flora 
regarded as semi-aquatic. 

A small percentage of the taxa have winged dissem- 
inules and were probably wind-dispersed, such as the 
Samaras of Cedrelospermum, Deviacer, Cruciptera, Pa- 
laeocarya, Palaeoplatycarya, and Pteronepelys, the 
tufted achenes of Platanus and Tanyoplatanus, and the 
seeds of Emmenopterys, Fimbrialata and Hydrangea. 
In extant forests, wind-dispersed species tend to best 
Tepresented among trees and lianas of the forest can- 
Ору, while bird-dispersed species are concentrated in 
the understory (Gentry, 1983). Gentry (1983) also not- 
ed that the percentage of species with wind-dispersed 
diaspores decreases as one proceeds from dry to wet 
forests in the neotropics. 

At least 80% of Nut Beds taxa appear to have been 
Well-suited for biotic dispersal. Nuts that commonly 
are dispersed by rodents today include Juglans, Cas- 
tanopsis and Quercus and it may be expected that Cor- 
Yloides, being similar to Corylus, was similarly adapt- 
ed. Although I have not observed obvious gnaw marks 
Оп any of the fossil nuts or seeds, it is likely that rodents 


played some role in seed dispersal. Among the mam- 
malian fauna of the Nut Beds, Orohippus, Hyrachyus 
and Telmatherium were browsers that may have par- 
ticipated in fruit dispersal. Birds were probably the 
most important dispersal agents for Nut Beds plants. 
As inferred from extant relatives and from morphology 
of the preserved fruit and seed remains, the over- 
whelming majority of Nut Beds species represent fleshy 
fruits, of the kind regularly dispersed by birds. Some 
examples include Actinidia, Anamirta, Annonaceae, 
Aphananthe, Celtis, Cornus, Ensete, Mastixia, Me- 
liosma, Nyssa, Prunus, Rhus, Sabal, Symplocos, Tap- 
iscia, Taxus, and the seven species of Vitaceae. 

Although seed size ranges greatly among the Nut 
Beds species, the occurrence of many species with large 
fruits and seeds is particularly striking (Scott, 1954; 
Tiffney, 1984). Based on a survey of seed size in Cre- 
taceous and Tertiary floras, Tiffney (1984) observed 
that most angiosperm seeds of the Cretaceous are small; 
large fruits first become common in the Early Tertiary. 
The abundance of large fruits in Eocene floras such as 
the Nut Beds probably coincides both with the radi- 
ation of mammal and bird taxa important as dispersal 
agents, and with the formation of closed canopy forest 
(Tiffney, 1984). A well-marked correlation between 
large seed size and establishment in shady, stable plant 
associations has been statistically documented in living 
tropical woody plants (Salisbury, 1942; Foster and Jan- 
son, 1985; Foster, 1986). In a closed canopy evergreen 
forest, large seeds have an adaptive advantage over 
smaller ones because they possess more food reserves 
for the embryo, enabling the seed to germinate and 
become established under subdued light conditions. 
Smaller seeds typically require more intense sunlight 
for sufficient photosynthesis for a seedling to become 
established, and are commonly produced in large 
quantities by plants that are successful at colonizing 
sun-lit gaps in the forest canopy. The high proportion 
of relatively large, biotically dispersed fruits and seeds 
in the Nut Beds assemblage may thus be an indication 
of closed canopy forest. 


CLIMATIC INTERPRETATION 


Fossil plants and animals of the Clarno Nut Beds 
reflect climatic conditions under which the biota ex- 
isted. The presence of crocodilian teeth and such ther- 
mophilic plants as cycads, palms (Sabal) and bananas 
(Ensete) gives an indication that the Nut Beds biota 
lived under typically frost-free, subtropical or tropical 
conditions. The Middle Eocene coastline was situated 
only about 100 km to the west (Nilsen and McKee, 
1979) so that the Clarno biota probably experienced 
the moderating climatic influence of the nearby ocean. 
Deposition of the Clarno Formation took place prior 


22 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


to the Neogene uplift ofthe Cascade and Coast Ranges 
that effected the present-day rain shadow. 

Paleoclimatic conditions are inferred by identifying 
floristic and/or physiognomic similarities of a fossil 
assemblage to a modern vegetation type of known cli- 
matic parameters. Discussion of paleoclimate requires 
a working definition of terms, because different inves- 
tigators have applied widely differing concepts of what 
constitutes such categories as tropical, subtropical, and 
temperate. For consistency with other paleobotanical 
literature Ihave adopted the definitions of Wolfe (1979) 
which are based on modern mean annual temperatures 
(MAT) for humid to mesic regions of eastern Asia: 
tropical >25°C; paratropical 20-25°С; subtropical 13- 
20°C; temperate 10-13°С; paratemperate 3-10?C; sub- 
temperate <3°C. These paramaters, along with mean 
cold month temperature and mean annual range of 
temperature, appproximately delimit distinct forest 
types within mesic vegetation (Wolfe, 1979). 

The high proportion oflianas already discussed (43% 
of the species with known familial affinities) supports 
previous interpretations that the Nut Beds flora rep- 
resents tropical or paratropical forest (Scott, 1954; 
Wolfe 1977), possibly with a multistratal canopy (Up- 
church and Wolfe, 1987). Observing that in Recent 
vegetation, “the great majority of woody lianas are 
restricted to tropical forests," Gentry (1992) computed 
the percentage of lianas per florula in temperate eastern 
North America at 6%, vs. 19% in continental tropical 
florulas. Gentry’s work did not specifically address li- 
ana diversity in subtropical or paratropical situations. 

Woods from the Nut Beds consistently show growth 
rings, indicating seasonality of temperature and/or pre- 
cipitation. However, true ring-porous wood structure, 
which typically is confined to temperate situations, has 
not been observed. Of the fruit and seed genera that 
have been identified to extant genera, at least 42% are 
deciduous-leaved today (Table 2). Fossil leaves from 
the Nut Beds are mostly of large size (67% of the di- 
cotyledonous leaf species exceed 10 cm in lamina 
length) and, at latest count (preliminary to a detailed 
work on the leaf flora), 33 of the dicot leaf species, or 
52%, are entire-margined. Additional research on leaves 
and woods of the flora may provide physiognomic data 
useful in testing the taxonomically-based assessment 
provided here. 

One approach to the assessment of paleoclimate is 
based on the growing conditions of extant taxa related 
to those identified in the fossil assemblage. The cli- 
matic preferences of present-day relatives of the fossil 
taxa are taken as an indication of the range of condi- 
tions under which the fossil flora might have existed. 
There are various problems inherent in using this uni- 
formitarian approach: 1) the climatic tolerances of a 


taxon may evolve to accommodate environmental 
change; 2) the current distribution of a taxon is con- 
trolled by many factors, only one of which is regional 
climate (other important factors include historic bar- 
riers to dispersal, and effects of interspecies competi- 
tion and pathogens); 3) any taxa that are misidentified 
may introduce erroneous climatic data. Because of 
problems that may result from mistaken identities, I 
have been particularly cautious in the systematic treat- 
ment of genera in this monograph, placing them in 
form- or stereotype genera, rather than in the “nearest 
modern equivalent” when characters diagnostic of a 
modern genus could not be demonstrated. Despite the 
limitations of taxonomically based assessments of pa- 
leoclimate, such analyses provide at least a coarse guide, 
and may be used to evaluate conclusions reached by 
other methods. 

Among the Nut Beds taxa that have been identified 
to Recent genera, most live today within the broad 
range of temperate to tropical vegetation (Table 2). 
Some of these, such as Ampelocissus, Anamirta, Di- 
ploclisia, Ensete, Iodes, Mastixia, Pyrenacantha, and 
Tinospora, are confined to tropical and paratropical 
vegetation today. Others are elements of temperate 
forest associations (Mixed Mesophytic and Mixed 
Broadleaved Deciduous forests, sensu Wolfe, 1979, as 
modified from Wang, 1961), and are not found in trop- 
ical vegetation, e.g., Cornus, Decodon, Emmenopterys, 
Fortunearites (inferred from extant Fortunearia and 
Sinowilsonia), Parthenocissus, Tapiscia, and Torreya. 
Table 2, listing extant genera of the Nut Beds flora and 
the vegetation types in which they occur today, shows 
the highest number of taxa (27 of 40 genera) occurring 
in the Mixed Mesophytic and Mixed Broadleaved Ev- 
ergreen forest communities, followed by the 24 genera 
with Paratropical Rain forest affinities, and 22 with 
Tropical Rain forest affinities. However, if one were 
to score the extinct genera of Menispermaceae and 
Icacinaceae as tropical, where these families predom- 
inate today, the larger proportion of Nut Beds taxa (37 
of 52 genera, or 71%) would be regarded as tropical to 
paratropical. 

Faced with a mixture of taxa, some that are exclu- 
sively tropical today and others that are exclusively 
temperate, interpretation of paleoclimate for the Nut 
Beds depends on the assumptions made and how the 
data are weighted. One might, for example, infer 4 
temperate environment for the Nut Beds and suggest 
that many elements have subsequently evolved into 
more tropical situations where they are found today. 
Or, alternatively, the Nut Beds may represent tropical 
or paratropical vegetation, some of the elements of 
which have since evolved to accommodate more tem- 
perate conditions. The second scenario is favored be- 


| 
| 
| 
| 


| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 23 


cause tropical to subtropical vegetation is also sug- 
gested by foliar physiognomic considerations (Wolfe, 
1977; Manchester, 1981). It is also possible that the 
apparent mixture of cool and warm elements is a gen- 
uine reflection of mixed sources, some of the cooler 
elements having been transported from higher eleva- 
tions. 

Presumably, the presence of frost intolerant taxa 
places more constraints on the inference of climate 
than the presence of frost tolerant genera. Although 
temperate taxa may survive in tropical situations, par- 
ticularly at higher elevations, tropical elements such 
as Ensete, a member of the banana family, would not 
be expected to persist in a setting with cold winters. 
Ensete occurs today in paratropical and tropical veg- 
etation of southeast Asia where the mean annual tem- 
perature falls within the ranges of 20-25°C (Paratrop- 
ical Rain forest, sensu Wolfe, 1979) and 25-28°C 
(Tropical Rain forest sensu Wolfe, 1979), and the mean 
annual range of temperature is low. The mean annual 
range of temperature varies from 2-19°C among dif- 
ferent weather stations in Paratropical Rain forest and 
2-11*C in Tropical Rain forest (Wolfe, 1979). The cold 
month mean temperature in these vegetation types is 
10°С or greater. If it is assumed that its climatic tol- 
егапсеѕ have not changed significantly since the Eo- 
cene, then the presence of Ensete suggests warm, moist, 
€quable climate, as is also consistent with the presence 
ОЁ cycads and palms. 

The Nut Beds provides just one example ofthe warm 
Vegetation that was widespread in the Eocene of the 
northern hemisphere (Wolfe, 1985). Tropical to par- 
atropical vegetation has also been recognized in the 
Eocene of southern Alaska (Wolfe, 1972, 1977) and in 
Europe (Reid and Chandler, 1933; Chandler, 1964). A 
marked temperature decline is apparent by the Early 
Oligocene, possibly corresponding to changes in ocean 
Currents effected by the Late Eocene separation of South 
America from Antarctica resulting in development of 
the circum-Antarctic current (Kennett, 1977; Parrish, 
1987) and the formation of deep water channels en- 
abling southward drainage of Arctic Ocean waters 
Caused by the rifting of Spitzbergen and Iceland from 
лш (McKenna, 1975; B. LePage, pers. comm., 

992). 


BIOGEOGRAPHIC CONSIDERATIONS 


The Nut Beds flora contributes a wealth of data use- 
ful in phytogeographic investigations. Floristic com- 
Parison, both with other Tertiary floras and with extant 
floras, provides insight into Ше spread of boreotropical 
forest in the Northern hemisphere. This section con- 
Siders the levels of taxonomic similarity of the Nut 
Beds flora with other fossil fruit and seed assemblages 


and with living floras. Comparison with other Tertiary 
floras indicates particularly close similarity with the 
Eocene flora of western Europe. Among modern-day 
vegetation, the closest taxonomic resemblance is found 
in eastern Asia. 


Comparison with other Fossil Floras 


Because of the high proportion of Nut Beds genera 
that are extinct and/or of indeterminate familial affin- 
ity, it is not possible to link each of the fossil fruit/ 
seed species with a corresponding leaf species. There- 
fore, the most informative paleofloristic comparisons 
are those made with other fossil fruit and seed assem- 
blages. Detailed comparison with other Paleogene leaf 
floras will become possible when the leaf assemblages 
of the Nut Beds and other Clarno localities are docu- 
mented. 


North American Fossil Floras 


The Late Eocene or Early Oligocene LaPorte flora 
of northern California, best known for its leaf assem- 
blage (Potbury, 1935), has recently yielded a good col- 
lection of well-preserved, lignitized fruits and seeds, 
many of which are shared at the generic, and possibly 
specific, levels with Nut Beds taxa. A preliminary tax- 
onomic list of LaPorte fruit and seed flora was pre- 
sented by Tiffney (in Doyle et al., 1988) along with an 
identification key and illustrations (those taxa indi- 
cated by asterisks (*) are shared with the Nut Beds 
flora): Dracontomelon type [probably Pentopercu- 
lum*], hamamelidaceous seed*, Quercus*, Pterocarya/ 
Cyclocarya type, Magnolia?*, Мазижа (Ganitrocera 
type)[probably Mastixioidiocarpum*], Odontocaryoi- 
dea type*, Stephania type [similar to Palaeosinomen- 
ium*], Tinospora type*, Nyssa*, Rhamnus?, Zanthox- 
ylum, Sapotaceae*, Tapiscia?*, Halesia?, Symplocos*, 
Vitis*, Parthenocissus*. It is noteworthy that the Nut 
Beds flora bears greater similarity to this younger flora 
than to other Middle Eocene fruit and seed assemblages 
in North America. 

The Middle Eocene Princeton chert flora of southern 
British Columbia, Canada includes a variety of ana- 
tomically preserved fruits and seeds that are revealed 
by sectioning and peeling the chert (Basinger and Roth- 
well, 1977; Cevallos-Ferriz and Stockey, 1988a, 1988b, 
1989, 1990, 1991). The flora is relatively low in di- 
versity, but includes several taxa in common with the 
Nut Beds flora, including Vitaceae (Cevallos-Ferriz and 
Stockey, 1989), Lauraceae, sabaloid Palmae (Erwin, 
1987), Decodon (Cevallos-Ferriz and Stockey, 19882), 
Prunus (Cevallos-Ferriz and Stockey, 1991), a mastix- 
ioid species similar to Mastixicarpum (Stockey, pers. 
comm., 1990), and the newly recognized Stockeycarpa 
(p. 111). The Princeton flora includes aquatic plants 


24 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


such as Nympheaceae (Cevallos-Ferriz and Stockey, 
1989), which are unknown from the Nut Beds, and 
appears to lack the diversity of lianas characteristic of 
the Nut Beds assemblage. Metasequoia, which is abun- 
dant in the Princeton flora (Basinger, 1981) is absent 
from the Clarno Formation, though it becomes abun- 
dant in the Lower Oligocene ofthe overlying John Day 
Formation (Manchester and Meyer, 1987). The full 
diversity of the Princeton flora remains to be docu- 
mented so I hesitate to provide statistical comparison 
between the Princeton and Nut Beds floras. However, 
the presence of several shared genera indicates that the 
geographic ranges of some of the Nut Beds taxa ex- 
tended into the northern part of the continent. 

Although it is known mostly from leaf impressions, 
the Middle Eocene (Lower Ravenian) assemblages of 
the Kushtaka and Kulthieth Formations in the Gulf 
of Alaska appear to be floristically very similar to the 
Nut Beds assemblage (Wolfe, 1972, 1977). Shared or 
similar genera include Alangium, Anamirta, Diplocli- 
sia, Limacia (cf. Davisicarpum), Mastixia, Meliosma, 
Palaeophytocrene, ?Pyrenacantha, Sabalites, and Vi- 
tis. These assemblages are similar to the Nut Beds flora 
in the high proportion of lianas. Geophysical and pa- 
leobotanical data suggest that the Gulf of Alaska floras 
are part of a tectonic block that was located farther 
south in the Eocene (Cowan, 1982; Saleeby, 1983; Ax- 
elrod, et al., 1991). 

In the southeastern United States, the Middle Eo- 
cene Claiborne flora of Tennessee and Kentucky is best 
known for fossil leaves (Berry, 1930; Dilcher, 1971), 
but has also yielded about 90 distinct taxa of fruits and 
seeds (Grote, 1989). Given the similarity in age, one 
might expect a high percentage of taxa to be shared 
between the Clarno and Claiborne floras. Although 
some of the same families are present, e.g., Annona- 
ceae, Hamamelidaceae, Juglandaceae, Leguminosae, 
Magnoliaceae, Sapotaceae, and Theaceae, generic sim- 
ilarity is minimal, including only Magnolia (Grote, 
1989), Nyssa (Dilcher and McQuade, 1967), Palaeo- 
carya (Dilcher, Potter and Crepet, 1976; Manchester, 
1987), and Sabal (represented by leaves, Daghlian, 
1978). Unlike the Nut Beds flora, the Claiborne in- 
cludes a large number oftaxa with present-day relatives 
in tropical America (Roth and Dilcher, 1979; Taylor, 
1990). The Clarno flora lacks the high diversity of 
legumes found in the Eocene of southeastern North 
America (Herendeen and Dilcher, 1990; Herendeen, 
1992). The apparent absence in the Claiborne flora of 
families that are diverse in the Clarno flora, such as 
Icacinaceae, Menispermaceae and Vitaceae, is con- 
spicuous. These differences suggests major floristic dis- 
continuity between northwestern and southeastern 
North America in the Middle Eocene. The floristic 
differences between the Clarno and Claiborne assem- 


blages might be due in part to climatic and edaphic 
factors, but may also reflect barriers to dispersal, in- 
cluding the prior mid continental sea of the Cretaceous 
and rise of the Rocky Mountains. High paleoelevations 
that have been inferred for the southern Rocky Moun- 
tains by the Late Eocene (Meyer, 1992) may have re- 
inforced floristic differentiation of western and eastern 
Eocene floras of North America. 

The Brandon Lignite flora of Vermont is relevant in 
this comparison as the only well-documented Tertiary 
megafossil flora of northeastern North America. Al- 
though not independently dated, the flora is thought 
to be Oligocene (Tiffney, 1981) or perhaps Early Mio- 
cene (Tiffney, 1985b). The Brandon flora appears to 
lack tropical elements such as Menispermaceae and 
Icacinaceae. Genera shared with the Nut Beds include 
Alangium (Eyde, Bartlett and Barghoorn, 1969), Mag- 
nolia (Tiffney, 1977), Vitis and Parthenocissus (Tiffney 
and Barghoorn, 1976), Nyssa (Eyde and Barghoorn, 
1963), and Symplocos (from pollen, Traverse, 1955; 
from endocarps, Tiffney, pers. comm., 1992). The 
composition and geographic placement of the Brandon 
flora provide support for the concept ofa Tertiary land 
connection across the North Atlantic (Tiffney, 1985b). 


European Fossil Floras 


Most of the families and many of the genera present 
in the Nut Beds are also known from the Eocene of 
Europe (Text-fig. 6). Comparable floras include the 
Early Eocene London Clay of southern England and 
the Middle Eocene Geiseltal and Messel floras of Ger- 
many (Table 3). The largest and most intensively in- 
vestigated of these, with about 200 described genera 
of fruits and seeds, is the London Clay (Reid and Chan- 
dler, 1933; Chandler, 1961b, 1964, 1978; Collinson, 
1983). The London Clay assemblage includes pyritized 
fruits and seeds collected from several localities of the 
London Clay Formation along the southern coast of 
England. The London Clay fossils were deposited in 
marine sediment, and include disseminules of man- 
grove vegetation and shark teeth that are absent in the 
Nut Beds. However, most of the London Clay plants 
are terrestrial, evidently deposited near the mouth of 
a large river, and provide a good record of regional 
vegetation. 

Similarity between the Clarno and London Clay flo- 
ras was recognized previously on the basis of prelim- 
inary investigation of the Nut Beds assemblage and 
has contributed to the concept of an Eocene boreo- 
tropical flora (Scott, 1954; Chandler, 1964; Wolfe, 1975; 
Tiffney, 1985b; Mai, 1989). It is now possible to quan- 
tify the level of similarity based on the entire Nut Beds 
fruit and seed collection. Of the 145 fruit and seed 
genera currently recognized from the Nut Beds assem- 
blage, 30 genera (2096) are shared with the London 


| 
| 
| 
| 
| 
| 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 25 


Clay flora (Table 3; Text-fig. 6). In addition, about 15 
Species (10%) are the same or are nearly indistinguish- 
able from those of the London Clay. These data cor- 
roborate other evidence for the existence of a land- 
connection between Europe and North America during 
the Early Tertiary (McKenna, 1975; Tiffney, 1985a, 
b). Dated as Early Eocene (Collinson and Hooker, 1987) 
the London Clay biota predates the Middle Eocene 
Nut Beds biota by about five million years. The timing, 
routes and rate of dispersal of individual taxa across 
land connection(s) between western Europe and west- 
ern North America are not well understood, but the 
younger age ofthe Nut Beds deposit indicates that the 
interchange of these taxa between Europe and North 
America must have occurred during or prior to the 
Middle Eocene. 

Fruit and seed floras of central Europe are not as 
diverse, or have not been as thoroughly documented, 
as the London Clay flora, but include additional taxa 
shared with the Clarno flora. The Middle Eocene Gei- 
seltal flora from extensive lignite deposits in eastern 
Germany includes well-preserved leaves (Rüffle, 1976), 
pollen (Krützsch, 1976) and fruits and seeds. Mai (1976) 
recognized 25 genera and 28 species of fruits and seeds 
from the Geiseltal, nine of which are shared with the 
Nut Beds: Castanopis, Iodes, Magnolia, Meliosma, 
Nyssa, Pinus, Sabal, Tapiscia, and Alangium. The 
Middle Eocene Messel flora in western Germany 
(Schaarschmidt, 1988) is a lacustrine deposit that in- 
cludes well-preserved compressions of leaves (Wilde, 
1989), and fruits and seeds (Collinson, 1982, 1988). 
About fifty genera of fruits and seeds are present in the 
Collections from Messel, about one quarter of which 
are shared with the Nut Beds. The shared genera in- 
Clude Cruciptera, Iodes, Magnolia, Mastixia, Melios- 
та, Palaeocarya, Palaeophytocrene, Palaeosinomen- 
ium, Pentoperculum, Prunus, Tapiscia, Tinospora, 
Vitis?, and Tripartisemen (Collinson, 1982, 1988 and 
Pers. obs.)?. Of these, Palaeocarya, Tripartisemen 
(“?Lythraceae” in Collinson, 1988) and Cruciptera 
(Manchester, 1991) are not recorded from the London 
Clay. 

Combining data from the London Clay, Messel and 
Geiseltal floras (Table 3), the percentage of Nut Beds 
genera shared with the Early to Middle Eocene of Eu- 
Tope is 24. Some of the Nut Beds taxa have not been 
Tecorded from the Early and Middle Eocene of Europe 
but are known from later Tertiary deposits. Examples 
include Actinidia (Kirchheimer, 1957; Friis, 1985), 
Cedrelospermum (Manchester, 1987b, 1989a)?, Celtis 
(Manchester, 1989c), Decodon (Friis, 1985), Hydran- 
Sea (Mai, 1985), Rhus (Friis, 1969), and Sabia (Geis- 
nn 


* Cedrelospermum is now also recognized at Messel. See Notes 
added in proof, p. 200. 


Total 


40 4 


London Clay 
30 4 


20 - 


Messel 


Number of Genera 


Geiseltal 


Text-figure 6.— Nut Beds fruit and seed genera shared with Eocene 
floras of Europe. Genera listed in Table 3. 


sert and Gregor, 1981). Most of these are no longer 
native to Europe, having succumbed to Plio-Pleisto- 
cene climatic cooling. 


Paleogeography 


The geographic routes of exchange between the Nut 
Beds flora and the London Clay, Messel and Geiseltal 
floras of western and central Europe are ofconsiderable 
phytogeographic interest (Tiffney, 1985a, b). Geo- 
physical data suggest that land bridges were open both 
across the North Pacific and across the North Atlantic 
during the Eocene (Parrish, 1987). If the connection 
was across Beringia, then one would expect to see 
marked floristic similarities with the Eocene of Asia. 
Although floristic ties with the Recent flora of eastern 
Asia are strong (discussed next section), the antiquity 
of this connection needs to be documented. In this 
regard, it would be desirable to make detailed com- 
parisons with floras of similar age in Asia. However, 
I am not aware of comparable Asian Eocene fruit and 
seed floras. Careful comparisons among leaf and pollen 
records are needed for a better understanding of the 
similarities and differences between North American 
and Asian Eocene floras, a project beyond the scope 
of the present study. 

Even if taxa were successful in spreading between 
North America and Asia via Beringia, the Turgai sea- 
way that separated Asia and Europe through at least 
the Early Eocene may have blocked exchange with Eu- 
rope (Tiffney, 1985b). Exchange across the North At- 
lantic via connections through Greenland and Iceland 
is viewed as more likely (McKenna, 1975; Tiffney, 
1985b), but Early Tertiary floras so far studied from 
Greenland and Spitzbergen give no clue to the presence 
of such thermophilic taxa as Menispermaceae and Ica- 
cinaceae. 


Comparison with Recent Floras 


The only genera of the Nut Beds flora that are still 
native to Oregon today are Pinus, Taxus, Celtis, Cor- 


26 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Table 3.—Clarno fruit and seed taxa shared with European Eocene floras.* 


Alangiaceae 


Alangium Le, Су 
Anacardiaceae 

Pentoperculum LC 
Annonaceae 

Anonaspermum Le 
Burseraceae 

Bursericarpum EG 
Cornaceae 

Cornus LC 

Langtonia Ee 

Mastixia LC, M 

Nyssa EGG 
Fagaceae 

Castanopsis G 
Flacourtiaceae 

Saxifragispermum LC 
Icacinaceae 

Тодез LC, G, M 

Palaeophytocrene LC, M 
Juglandaceae 

Cruciptera M 

Palaeocarya ECE M 
Lauraceae 

Laurocalyx EC 

Laurocarpum EC 
Magnoliaceae 

Magnolia LC, G, M 


Menispermaceae 

Atriaecarpum ще: 

Davisicarpum BG 

Diploclisia LC 

Eohypserpa ще 

Palaeosinomenium LC, M 

Tinospora LC, M 

Tinomiscoidea LC 
Palmae 

Sabal е 76, 
Rosaceae 

Prunus M 
Sabiaceae 

Meliosma LC, б, М 
Sapindaceae 

Palaeoallophylus ще 
Staphyleaceae 

Tapiscia LC, G,M 
Symplocaceae 

Symplocos LC 
Theaceae 

Cleyera TG 
Vitaceae 

Vitis LC, M 

Ampelopsis LC 

Parthenocissus LC 
Incertae Sedis 

Carpolithus bellispermus LC 

Tripartisemen M 


LC = London Clay; G = Geiseltal; M = Messel. * See also Notes added in proof, p. 200. 


nus, and Quercus. The majority of extant genera of the 
Nut Beds flora are found today in other geographic 
regions. Table 4 and Text-fig. 7 show the distribution 
of Recent genera known from the Nut Beds in selected 
modern phytogeographic provinces (southeast Asia, 
Malesia, western North America, Central America, 
South America, Europe and Africa). Strongest ties are 
with Southeast Asia and Malesia. 

Within the Americas, the closest floristic ties are 
with eastern North America and with central America. 
Twenty-three genera, or 56% of the extant genera known 
from the Nut Beds, occur in eastern North America. 
Although most of these occur in other continents as 
well, Calycocarpum and Decodon are endemic to east- 
ern North America. Twenty-two genera, or 54%, live 
today in Central America. However, none of these are 
presently endemic to the region. Sabal, distributed in 
the southeastern U.S. and tropical America, is one of 
the few genera of the Clarno assemblage that is limited 
to the New World in its present-day distribution. The 
genus was formerly more widespread, however, with 
Eocene records in Europe (e.g., Mai, 1976) and Asia 
(Huzioka and Takahasi, 1970). 

Twenty-four percent of the Nut Beds genera occur 


today in Europe, although none are endemic there. 
This percentage, although significantly less than those 
computed for the present-day floras of southeastern 
Asia, Malesia, southeastern North America and central 
America (Table 4), matches that presented above in 
comparing the Nut Beds flora with Early to Middle 
Eocene floras of Europe (24%). However the similarity 
of these values for Eocene and Recent flora of Europe 
probably is an artifact of the quantitative analysis. All 
152 genera, both extinct and extant, are included in 
the base number for comparing fossil assemblages, 
whereas only the 41 extant genera form the basis of 
comparison with the Recent flora. It is important to 
note that although 36 of the Nut Beds genera are shared 
with the Eocene flora of Europe, only ten are found in 
Europe today. Many of the extant genera shared be- 
tween the Clarno and the London Clay floras are now 
absent from Europe and are disjunct between Asia and 
eastern North America and/or central America, for 
example, Nyssa, Hydrangea, Symplocos; and others 
are endemic to southeastern Asia. 

The strongest geographic affinities are with south- 
eastern Asia (Indochinese Region, sensu Takhtajan, 
1986) and Malesia (Malesian Region, sensu Takhtajan, 


| 
| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 27 


Table 4.— Present geographic distribution of extant genera rep- 
resented in the Nut Beds flora. 


genus seAs Mal wNA eNA cAm sAm Eur Afr 

Actinidia x x 
Alangium x x x 
Ampelocissus х 7 x ? 
Ampelopsis x ? x 
Anamirta x x 
Aphananthe x x х х 
Bumelia х x х 
Calycocarpum x 
Castanopsis х x у 
Celtis x x x x x x x x 
Cleyera x x 
Cornus x x x x x x x x 
Decodon x 
Diploclisia x x 
Emmenopterys x 
Ensete x x x 
Hydrangea x x x x 
Zodes x X х 
Juglans E x x x x xe 
Lindera х Y x 
Magnolia x X x x 
Mastixia х х 
Мейозта x x x x 
Nyssa x x x x 
Pathenocissus x х x х 
Pinus x х х x x в х х 
Platanus x x х * х 
Prunus х x х x x х 
Pyrenacantha x x x 
Quercus x x x x x x x 
Rhus x x x x x x x x 
Sabal x х х 
Sabia x Ж 
Schisandra x x x 
Symplocos x х x x x x 
Tapiscia x 
Taxus x x х x x x 
Tinospora x x x 
Tor reya x i x x 
Trema x X x x x 
Vitis x x х х х x 

Totals 97 25 14 25 22 12 10 13 


SeAs = Southeast Asia; Mal = Malesia; eNA = eastern North Amer- 
lca; sAm = South America; Eur = Europe; Afr = Africa. 


1986). Among Recent floras, the highest number of 
genera shared with the Nut Beds assemblage is found 
In eastern Asia. Thirty-seven genera, or 9096, are native 
to southeastern Asia. The flora of Malesia shares 25 
genera with the Nut Beds. Genera that are exclusively 
Asian or Malesian today include Actinidia, Mastixia, 
Anamirta, Sabia, Tapiscia. Although Meliosma occurs 
In central and South America as well as in Asia, four 
Of the Nut Beds species conform most closely to sub- 
genus Meliosma, section Meliosma, which is exclu- 
Sively oriental in its Recent distribution (Van Beuse- 
kom, 1971). The close floristic similarity with 
Southeastern Asia and Malesia probably reflects the 


Number of Genera 


Region 


Text-figure 7. — Present geographic distribution of Extant Nut Beds 
genera. Abbreviations and data from Table 4. 


status of this area as a refugium for once-widespread 
thermophilic genera that could not withstand the ef- 
fects of climatic cooling and glaciation at the end of 
the Tertiary in other parts of the northern hemisphere. 


INTRODUCTION TO SYSTEMATIC 
PALEONTOLOGY 


New species and genera have been named according 
to procedures required by the International Code of 
Botanical Nomenclature (Greuter, 1988). In order for 
a new species to be validly published, the new name 
must be accompanied by a description or diagnosis 
(art. 32.1). Fossil plants are excluded from the require- 
ment that the diagnosis be written in Latin (art. 36.1). 
A new genus and species may be created, and the names 
of a genus and a species may be simultaneously vali- 
dated, by provision ofa single description or diagnosis 
(art. 42.1). I have therefore given a combined generic 
and specific description for the many new, currently 
monotypic, genera presented here. Type and cited 
specimens are housed in the indicated museum col- 
lections. Specimens are cited as holotypes, paratypes, 
lectotypes and syntypes as defined in the Botanical 
Code of Nomenclature (Greuter, 1988). In addition, 
specimens that are not a part ofthe original description 
of species, but which are helpful in understanding the 
species may be designated hypotypes, as defined pre- 
viously in zoological (Schenk and McMasters, 1956) 
and paleobotanical (e.g., MacGinitie, 1953, p. 79) lit- 
erature. 

In naming species of fruits and seeds in this work, 
Ihave tended toward conservatism, and have probably 
erred more on the part of lumping than splitting. By 
contrast, I consider that the classic monographs on the 
London Clay flora (e.g., Reid and Chandler, 1933; 
Chandler, 1978) reflect a greater tendancy for taxo- 
nomic splitting. An important objective has been to 
document the major taxonomic groups represented, 


28 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


thereby highlighting taxa available to the research com- 
munity for more detailed individual systematic inves- 
tigations. Many of the extant families and genera rec- 
ognized have never been analyzed by cladistic 
methodology, and may be in need of revision. In at- 
tempting to classify the entire fossil flora, however, it 
has been necessary to accept the taxonomic treatments 
currently available, rather than attempting to revise 
intrafamilial classifications. 

Species are assigned to a modern genus only when 
sufficient diagnostic characters are present to rule out 
affınities with other extant genera and/or families. Oth- 
er species are assigned to extinct, form, or stereotype 
genera according to the criteria set out on pp. 17-18. 

In paleobotanical literature, the generic name Car- 
polithus Schlotheim traditionally has been used as a 
catchall for fossil fruits and seeds of unknown system- 
atic affinities (Andrews, 1970). Different "species" of 
Carpolithus are often so different from one another that 
they probably represent different biological genera and 
families. I have given unique generic names to most 
of the form genera described from the Nut Beds but 
have placed an additional 21 taxa as unnamed species 
of Carpolithus. Thus, in comparing the generic diver- 
sity of different fossil localities, it is important to be 
mindful ofthe hidden diversity lurking under the name 
Carpolithus. It should also be noted that different spe- 
cies assigned to stereotype genera such as Laurocarpum 
and Anonaspermum may in reality represent different 
genera, but information from other plant organs would 
be required to determine the “real” generic diversity 
represented by such taxa. 

Although leaves and woods co-occur with the fruits 
and seeds at the Nut Beds locality, no actual attach- 
ments have been found, and the high floral diversity 
combined with the large allochthonous component of 
the flora makes it difficult or impossible to determine 
which of the woods, leaves and reproductive structures 
might be conspecific. Therefore, I have refrained from 
assigning the same epithet to different isolated organs. 
It is conceptually intriguing to reconstruct fossil “whole- 
plant species” from their isolated organs; such recon- 
structions cannot be proven correct, however, unless 
there are specimens preserving direct attachment be- 
tween the organs. In some instances evidence of co- 
occurrence at numerous localities and shared modern 
familial affinity of isolated organs provides good evi- 
dence to infer that particular species of leaves and re- 
productive structures were perhaps produced by the 
same biological species, as in the case of the Clarno 
plane tree (Manchester, 1986). In the discussion ac- 
companying each family of fruit or seed recognized 
here, I indicate known records based on other organs 


from the Clarno Formation, recognizing that some of 
them could in reality represent the same biological 
species. 


ORGANIZATION 


The descriptive systematic section is organized into 
two parts: The first part includes genera that can be 
confidently assigned to extant plant families. These are 
arranged alphabetically by family, treating the gym- 
nosperms first, followed by the angiosperms. The sec- 
ond part treats genera of uncertain familial affinity, 
ordered alphabetically, followed by a numerical se- 
quence of species informally attributed to Carpolithus. 
Families have been organized alphabetically, rather 
than phylogenetically, because there is more agreement 
among botanists as to the sequence of the alphabet 
than to the best phylogenetic arrangement. Phyloge- 
netic schemes are subject to continual change and re- 
finement and, while clearly important in expressing 
current views of relationships, are cumbersome and 
unstable as indexing systems. Table 5 presents a phy- 
logenetically arranged list of the Nut Beds fruit and 
seed genera following, with some modification, the ar- 
rangement of Cronquist (1981). Unless otherwise in- 
dicated, the numbers for species diversity and the geo- 
graphic ranges of extant genera were obtained from 
Mabberley, 1989. 


TERMINOLOGY 


Geometric symmetry, in particular reflectional point 
symmetry (Lockwood and MacMillan, 1978), has 
proven very useful in describing the three-dimensional 
shapes encountered among fruits and seeds. An object 
is said to be bilaterally symmetrical if there is a single 
plane of mirror symmetry. Following the same logic, 
I have used the term quadrilaterally symmetrical when 
there are two planes of mirror symmetry. These two 
planes usually intersect at right angles along the long 
axis of the fruit. 

In describing the fossil seeds, I have generally used 
the term seed coat, rather than testa or tegmen, because 
the terms testa and tegmen have been used in a re- 
stricted sense that implies knowledge of ontogeny (Cor- 
ner, 1976; Schmid, 1986). Seed coat is a neutral de- 
scriptive term for the wall of the seed, whereas the 
terms tegmen and testa (sensu Corner, 1976) require 
an interpretation that is better left to discussion. Like- 
wise, for fruits, I have applied the term endocarp in а 
loose sense to refer to the hard, inner layer of the fruit, 
regardless of its ontogenetic derivation. It is this hard 
layer of the pericarp (fruit wall) that is most often 
preserved in the fossil specimens, although occasion- 
ally the meso- and exocarp are preserved. Cell layers 


| 
| 
| 
| 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 29 


Table 5.—Systematic list of Nut Beds genera with known modern familial affinities. Angiosperms listed in phylogenetic sequence similar 


to that of Cronquist (1981). 


Gymnospermae 
Order Coniferales 
Family Taxaceae 
Genus Taxus 
Torreya 
Diploporus 
Family Pinaceae 
Genus Pinus 
Angiosperms 
Class Dicotyledonae 
Subclass Magnolididae 
Order Magnoliales 
Family Magnoliaceae 
Genus Magnolia, 3 spp. 
Family Annonaceae 
Genus Anonaspermum, 3 spp. 
Order Laurales 
Family Lauraceae 
Genus Lindera 
Laurocalyx 
Laurocarpum, 3 spp. 
Order Illiciales 
Family Schisandraceae 
Genus Schisandra 
Order Ranunculales 
Family Menispermaceae 
Genus Anamirta 
Atriaecarpum 
Calycocarpum 
Chandlera 
Curvitinospora 
Davisicarpum 
Diploclisia 
Eohypserpa 
Odontocaryoidea 
Palaeosinomenium 
Tinospora, 2 spp. 
Tinomiscoidea 
Thanikaimonia 
Family Sabiaceae 
Genus Meliosma, 5 spp. 
Sabia 
Subclass Hamamelidae 
Order Hamamelidales 
Family Platanaceae 
Genus Macginicarpa 
Platanus 
Tanyoplatanus 
Family Hamamelidaceae 
Genus Fortunearites 
Order Urticales 
Family Ulmaceae 
Genus Aphananthe 
Celtis, 2 spp. 
Trema 
Cedrelospermum 
Order Juglandales 
Family Juglandaceae 
Genus Juglans 
Cruciptera 
cf. Palaeocarya 
Paleoplatycarya? 
Order Fagales 
Family Fagaceae 
Genus Quercus 
Castanopsis 
Family Betulaceae 
Genus Coryloides 
Kardiasperma 
Subclass Dilleniidae 
Order Theales 
Family Actinidiaceae 
Genus Actinidia 


Family Theaceae 
Genus Cleyera 
Order Violales 
Family Flacourtiaceae 
Genus Saxifragispermum 
Order Ebenales 
Family Sapotaceae 
Genus Bumelia?, 2 spp. 
Family Symplocaceae 
Genus Symplocos 
Subclass Rosidae 
Order Rosales 
Family Hydrangeaceae 
Genus Hydrangea 
Family Rosaceae 
Genus Prunus, 2 spp. 
Order Fabales 
Family Leguminosae 
Genus Leguminocarpon 
Order Myrtales 
Family Lythraceae 
Genus Decodon 
Order Cornales 
Family Alangiaceae 
Genus Alangium 
Family Cornaceae 
Genus Cornus 
Langtonia 
Mastixia 
Mastixioidiocarpum 
Mastixicarpum 
Nyssa, 3 spp. 
Order Celastrales 
Family Icacinaceae 
Genus Paleophytocrene, 2 spp. 
Iodes, 2 spp. 
Pyrenacantha 
Comicilabium 
lodicarpa, 2 spp. 
Order Rhamnales 
Family Vitaceae 
Genus Ampelocissus, 2 spp. 
Ampelopsis 
Parthenocissus, 2 spp. 
Vitis, 2 spp. 
Order Sapindales 
Family Staphyleaceae 
Genus Tapiscia 
Family Sapindaceae 
Genus Palaeoallophylus, 2 spp. 
Deviacer 
Family Burseraceae 
Genus Bursericarpum, 2 spp. 
Family Anacardiaceae 
Genus Pentoperculum 
Rhus 
Order Apiales 
Family Araliaceae 
Genus Paleopanax 
Subclass Asteridae 
Order Rubiales 
Family Rubiaceae 
Genus Emmenopterys 
Monocotyledons 
Subclass Arecidae 
Order Arecales 
Family Palmae 
Genus Sabal, 2 spp. 
Subclass Zingiberidae 
Order Zingiberales 
Family Musaceae 
Genus Ensete 


30 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


may be described as uni-, bi-, or multiseriate, indi- 
cating the thickness of the layer in numbers of cells. 
The orientation of cells relative to the seed or fruit 
surface morphology may also be important. Cells are 
oriented anticlinally, if their long axes are perpendic- 
ular to the trend of the adjacent external or internal 
surface. Cells are oriented periclinally if their long axes 
are parallel to the trend of the adjacent external or 
internal surface. Periclinally oriented cells may at the 
same time be horizontally or vertically oriented with 
respect to the principal axis of the fruit or seed. 


MEASUREMENTS 


If a species is known from three or fewer specimens, 
then the individual measurements of each are listed in 
the description. If more than three are measured, then 
the range, mean (avg.) and standard deviation (SD) are 
presented followed by the number of specimens mea- 
sured (n). Broken specimens vary in completeness and 
sometimes provide only one or two of the three spatial 
dimensions (length, width and/or thickness), hence, the 
number of measurement indicated for length may dif- 
fer from those given for width and/or thickness. 


ABBREVIATIONS 


Specimen numbers are prefixed by the appropriate 
institutional abbreviation, indicating where the spec- 
imen is deposited. The largest collections are at the US 
National Museum (USNM), Washington D.C. and the 
Florida Museum of Natural History, Gainesville (UF). 
Additional smaller collections were studied at the Uni- 
versity of California Museum of Paleontology, Berke- 
ley, California (UCMP), the University of Michigan 
Museum of Paleontology, Ann Arbor, Michigan (UM), 
Harvard University, Cambridge, Massachusetts (HU), 
the Burke Memorial Washington State Museum, Uni- 
versity of Washington, Seattle (UWBM), and the Or- 
egon Museum of Science and Industry, Portland 
(OMSI). Reference is also made to fossils from other 
localities deposited at the Natural History Museum, 
London (BM) and the Peabody Museum of Yale Uni- 
versity, New Haven (YPM) and to extant seeds from 
the National Seed Herbarium of the Beltsville Agri- 
cultural Research Center, Beltsville, Maryland (BARC), 
the herbaria of the Smithsonian Institution, Washing- 
ton, DC (US), Missouri Botanical Garden (MO), Rijk- 
sherbarium, Leiden (L), University of Florida (FLAS) 
and the Arnold (A) and Gray (GH) herbaria of Harvard 
University, Cambridge, Massachusetts. 

The authors’ names for families, genera and species 
are presented at the first occurrence in the text, and 
may be abbreviated. Standard abbreviations for the 


names of botanists are presented in Mabberley (1989, 
pp. 661—706). 

In the plate captions, the abbreviations preceding 
the magnification refer to techniques used in photog- 
raphy. “РС”, or palladium-coated, is indicated for 
specimens that were coated to enhance reflected light 
microscopy (see materials and methods). “TL” refers 
to transmitted light microscopy, and ““RL” to reflected 
light micrography, while “SEM” denotes scanning 
electron microscopy. When not specified, specimens 
were photographed by reflected light, generally with 
the light source from the upper left and lower right. 


Family PINACEAE Lindley 
Genus PINUS L. 


Pinus sp. 
Plate 1, figures 6-9 


Description.— Pollen cone elongate, with central axis 
1.2 mm thick surrounded by numerous helically ar- 
ranged sporophylls, about 30 sporophylls per cycle; 
cone width 5.2 mm, length more than 12 mm (speci- 
men incomplete); lower part of cone with a bulge formed 
by large curved scales 4.5-5.5 mm long; sporophylis 
1.5-1.8 mm long, thickened and upcurved distally; 
pollen sacs paired, abaxial on each sporophyll, 0.7-0.8 
mm long. 

Specimen.— UF 9334. 

Discussion.— Although no seeds of Pinus have been 
identified from the Nut Beds, the genus is represented 
by wood fragments, dispersed pollen, and by a pollen 
cone. The pollen cone, described above, is known from 
a single specimen preserved in siltstone from the upper 
part of Face 1 of the Nut Beds (Pl. 1, fig. 6). Sectioned 
longitudinally (Pl. 1, figs. 7-9), the specimen shows the 
arrangement of microsporophylls typical of Pinaceae. 
The pollen itself is poorly preserved, possibly imma- 
ture. Similar pollen cones (Pl. 1, fig. 3), preserved as 
compressions, occur in association with foliage of a 
5-needle pine and seeds of the articulate type (Pl. 1, 
figs. 4, 5) at the West Branch Creek locality of the 
Clarno Formation. Although the single specimen from 
the Nut Beds is broken so that the full length cannot 
be determined, comparison with the West Branch Creek 
specimens indicates a length of about 4 cm. Abraded 
pinaceous seed cones also occur in the Nut Beds (е.Р., 
Pl. 1, fig. 2), but because of poor preservation, they 
have not been identified to genus. 


Family TAXACEAE Gray 


The Taxaceae, or yew family, has five extant genera 
and about 20 species of trees and shrubs (Price, 1989). 
Although the family is well documented by foliage of 


| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 31 


various genera in the Tertiary of Europe and North 
America, reports based on seeds are rare. Three genera 
of taxaceous seeds occur in the Nut Beds: Taxus, Tor- 
reya, and a new extinct genus, Diploporus. A single 
taxaceous leaf, with cuticle corresponding to that of 
Torreya, was recovered from the White Cliffs locality 
of the Clarno Formation. The leaf is noteworthy for 
its large size: 70 mm long, 7 mm wide. 


Genus TAXUS L. 


Taxus masonii sp. nov. 
Plate 2, figures 1-11 


Etymology.—This species is named for Herbert L. 
Mason, in recognition of his pioneering investigations 
of fossil conifers from western North America. 

Description.—Seed bilaterally symmetrical, lentic- 
ular in cross-section, nearly elliptical in face view, base 
truncate, apex keeled; length 4.3-6.2, avg. 5.3 mm 
(SD=0.53, n=10), width across plane of symmetry 4.5— 
5.6, avg. 4.9 mm (SD=0.34, n=10), thickness 3.3-4.0, 
avg. 3.7 mm (SD=0.28, n=10), length/width ratio 1.0- 
1.1; basal truncation perpendicular to central axis of 
Seed, elliptical with a pair of small circular vascular 
Scars; apical keel slightly arched as viewed from above, 
with a central micropylar scar; seed coat about 250- 
300 um thick, composed of isodiametric cells 25-60 
um in diameter; two vascular strands, one along each 
Side in the dorsiventral plane, arising from the basal 
truncation and passing into the seed cavity in the lower 
% of the seed. 

Specimens.— Holotype: USNM 355474. Paratypes: 
UCMP 10622, UF 6643, 6644, 9474—9478, 9801,9802, 
USNM 354997, 354998, 355000, 355406. 

. Discussion.— Taxus is represented by permineral- 
12е4 seeds from the Nut Beds that are readily recog- 
nized by their lensoid cross section, keeled apex and 
truncate base (Pl. 2, figs. 1-8). The pointed end rep- 
Iesents the micropyle, and the basal truncation cor- 
Tesponds to attachment of the aril with the seed. The 
Pair of vascular scars on the basal truncation (РИ 2 
figs, 7, 8) are diagnostic for the genus. Although extant 

ахиѕ seeds may vary from two- to three-fold sym- 
Metry, these fossil seeds are nearly all bilaterally sym- 
Metrical. The morphology and vascular supply of the 
Seed, as seen in serial transverse sections (Pl. 2, figs. 
9-11), is very similar to that observed in extant species 
(Dupler, 1920). 

Taxus grows today in central Malesia and Mexico 
45 well as in north temperate areas. The genus has also 
been confirmed from the Oligocene and Miocene of 
eastern Europe through careful study of cuticularly pre- 
Served leaves (Куасек, 1982). 


Genus TORREYA Arn. 


Torreya clarnensis sp. nov. 
Plate 2, figures 12-16; Text-figure 8A 


Etymology.— The epithet clarnensis refers to the 
Clarno Formation, source of the specimens. 

Description.—Seed bilaterally symmetrical, obovate 
in face view, fusiform in lateral view, ellipticalin trans- 
verse section; length 19.5-20.8 mm, width 10.7-11.3 
mm, thickness across the dorsiventral plane 9.5-9.6 
mm, length/width ratio 1.7-1.9; base acute-rounded, 
apex rounded 1n face view, acute in lateral view, with 
the dorsal and ventral faces meeting to form a keel in 
the apical % of the seed; micropylar scar at the mid- 
point of the apical crest; two circular vascular scars 
situated in the dorsiventral plane, one on either side 
of the seed; the vascular scars positioned at the level 
of greatest seed width, about Y of the length from the 
apex. 

Specimens.— Holotype: UF 6510. Paratypes: USNM 
353966, 422380. 

Discussion.— Torreya clarnensis is represented only 
by a few specimens but they present enough characters 
to establish firmly the affinities with extant Torreya. 
The apical keel, the pair of pores (vascular scars) pierc- 
ing the seed coat near the apex in the dorsiventral plane 
(Pl. 2, fig. 13), and the narrowed, acute-rounded base 
are characters diagnostic of the extant genus. The pair 
of vascular pores are closer to the apex (Text-fig. 8A) 
than they are in Diploporus (Text-fig. 8D), but not as 
close to the apex as in the extant species of Torreya 
(Text-figs. 8B, C). 

Torreya is a genus of trees with five extant species 
distributed in North America and eastern Asia. The 
North American species are restricted to a few sites in 
California and northwestern Florida. The Asian spe- 
cies are limited to an east-west belt extending from 
Honshu over the Yang-tse valley to eastern Burma. 
The genus has also been recognized on the basis of 
upon fossil foliage from the Late Eocene Florissant and 
Oligocene Bridge Creek floras (MacGinitie, 1953), and 
from Oligocene and Miocene localities Europe (Kva- 
сек, 1982). E 


Genus Diploporus gen. nov. 


Etymology.—Diplo (Gr = twofold) + poros (Gr = 
hole, passage) referring to the pair of vascular scars in 
the seed coat. 

Generic diagnosis.—Seed subovoid, base rounded, 
apex wedge-shaped; apical keel slightly arched as viewed 
from above, with a central micropylar scar; seed coat 


32 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Text-figure 8.—Seeds of Taxaceae in face view and lateral view, showing apical keel and variable position of the pair of vascular pores in 
the seed coat. A, Torreya clarnensis sp. nov., with pores positioned about of seed length below apex. B, T. grandis Fort. from Kuling, 
China, showing pointed base, apical pores. C, T. taxifolia Arn. from northwestern Florida showing apical pores. D, Diploporus torreyoides 
sp. nov., showing apical keel, pair of pores in the lower half, rounded base. Scale bar = | cm. 


pierced by a pair of circular vascular scars, one on each 
side in the dorsiventral plane at, or slightly below, the 
equator. 

Type species.— Diploporus torreyoides sp. nov. 


. Diploporus torreyoides sp. nov. 
Plate 3, figures 1-16; Text-figure 8D 


Etymology.— The epithet refers to similarity with 
seeds of extant Torreya. 

Description.—Seed bilaterally symmetrical, subo- 
void, ovate to circular in face view, pyriform in lateral 
view, wide-elliptical in transverse section; length 6.7- 
8.0, avg. 7.9 mm (SD=0.5, n=25), width 5.4-8.0, avg. 
6.9 mm (SD=0.6, n=25), thickness 3.6-7.0, avg. 5.8 
mm (SD=0.78, n=25), length/width ratio 1.0-1.3; base 


rounded; apex rounded in face view, angled in lateral 
view, with the dorsal and ventral faces meeting to form 
a sharp crest in the apical У of the seed with a central 
micropylar scar; two circular vascular pores situated 
in the dorsiventral plane, one on either side ofthe seed, 
positioned at the level of greatest seed width, about Y 
to % of the length from the apex; aril preserved in some 
specimens, completely covering seed, 1.0-1.5 mm thick, 
outer surface of aril with numerous short longitudinal 
grooves; seed coat smooth; embryo straight, 1.0 mm 
in diameter, 6.4 mm long, aligned with the central axis 
of the seed. 

Specimens.— Holotype: UF 8542. Paratypes: UE 
8501-8541, USNM 355072 (2 specimens), 355471, 
355479, 355481 (9 specimens), 355489 (17 speci- 
mens), 424744, 424745, 435098, OMSI Pb217, 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 33 


Pb1348, internal casts; UF 6531, 9587, 9803, 9804, 
USNM 355569, 446078, specimens with intact seed 
coat and aril. 

Discussion.—Represented by more than 75 speci- 
mens, this species is among the more common fossils 
from the Nut Beds, and is particularly abundant near 
the top of Face 3. Most specimens are smooth-sur- 
faced, representing internal molds of the seed coat (Pl. 
3, figs. 9-12); a few possess an additional outer layer 
about one mm thick with a wrinkly external surface 
(Pl. 3, figs. 1-8). This outer layer is poorly preserved 
anatomically but apparently represents the aril. Salient 
features of these fossils are the rounded base, keeled 
apex and the pair of conspicuous pores, one on either 
side in the dorsiventral plane (Pl. 3, figs. 6, 8, 11—13). 
Longitudinal and transverse sections reveal a slender 
cylindrical embryo (Pl. 3, figs. 14-16) that is medial 
within the seed. 

Diploporus is interpreted as an extinct genus of Tax- 
aceae based upon several important similarities with 
Taxus and Torreya, including the pair of vascular bun- 
dles, the central linear embryo and the apical keel that 
is slightly arched as viewed from above, with a central 
micropylar scar. The seed base is rounded, however — 
Dot truncate as in Taxus, and not pointed as in Torreya. 
The tissue surrounding the embryo within the seed 
Cavity is cellularly preserved in many of the specimens, 
and has a speckled appearance (Pl. 3, figs. 14-16) like 
that of Torreya. Diploporus seeds are larger than those 
of Taxus and smaller than those of Torreya. The pair 
of vascular pores in the dorsiventral plane corresponds 
to those seen in Torreya, but in the fossil these scars 
are positioned at or slightly below the equator, rather 
than near the apex (Text-fig. 8). 

Seeds of this morphology have not, to my knowl- 
edge, been described from the Tertiary before, and it 
is of some interest to confirm the presence of extinct 
Taxaceae surviving into the Tertiary. As this manu- 
Script was going to press, I noticed an apparent close 
Similarity of this genus to the Cretaceous genus Vesquia 
Bertrand from the Wealden Formation of Belgium (Al- 
Vin, 1960). Although I have not seen specimens of 
Vesquia, the genus was described in good detail and 
illustrated with line drawings (on the basis of two well 
Preserved liginfied seeds) by Alvin (1960). Vesquia seeds 
are similar in size and shape to those Diploporus, with 
а corresponding pair of pores (“two formamina") in 
the lower half. However, the specimens of Vesquia are 
More globose and appear to lack an apical crest. In 
addition, the basal surface of the Vesquia seeds have 
4 pair of grooves leading to the foramina, which are 
Not seen in Diploporus. Vesquia is also considered to 
belong to the Taxaceae and, among extant genera, com- 


pares most closely to Torreya (Alvin, 1960). It is pos- 
sible that additional work may reveal that the Clarno 
taxon is a hold-over from the Mesozoic. Indeed, both 
of the living genera, Torreya and Taxus, are believed 
to extend back to the Jurassic (Florin, 1958; Harris 
1976). 


Family ACTINIDIACEAE Hutch. 


This family includes four modern genera: Clema- 
toclethra, Sladenia, Saurauria and Actinidia. The lat- 
ter, which provides the kiwi fruit of commerce, is the 
best known. Saurauria has been reported on the basis 
of leaves from the middle Eocene of Alaska (Wolfe, 
1977) and Wyoming (MacGinitie, 1974). Seeds as- 
signed to the fossil genus Pasternackia (described, p. 
104) are similar to those of Saurauria, although affin- 
ities remain uncertain. The following constitutes the 
first recognition of Actinidia from the North American 
fossil record. 


Genus ACTINIDIA Lindl. 


Actinidia oregonensis sp. nov. 
Plate 4, figures 1—4 


Etymology.—The epithet refers to the state of Ore- 
gon. 

Description.—Seed bilaterally symmetrical, anatro- 
pous, elongate-elliptic in face view, laterally flattened, 
keeled in the plane of symmetry, lensoid in cross sec- 
tion; length 3.2-4.4, avg. 3.8 mm (SD=0.46, n=5), 
width 2.2-3.0, avg. 2.5 mm (SD=0.29, n=7), thickness 
1.0-1.1, avg. 1.1 mm (SD=0.05, n=7); base with ob- 
tusely pointed micropyle and laterally adjacent hilum, 
apex rounded; seed coat ca. 0.1 mm thick, surface re- 
ticulate, made up of thick-walled, polygonal (five- to 
seven-sided), isodiametric cells 190 to 240 um de- 
creasing in size toward the raphe and antiraphal edges 
of the seed; 15—20 cells wide in face view. 

Specimens.— Holotype: UF 6292. Paratypes: UF 
6293, 6294, 6498, 8580, 9218, USNM 355370. 

Discussion. — This species is represented by chalced- 
ony seed casts. The distinctive reticulate surface of 
these fossil seeds (Pl. 4, figs. 1-4) matches that in extant 
Actinidia seeds, where the outer layer of the seed coat 
is formed by polygonal cells with strongly thickened 
lateral walls. This surface ornamentation, combined 
with seed shape, and position of the hilum and micro- 
pyle, confirms the identification of these fossils as Ac- 
tinidia. 

Actinidia is a genus of about 30 species of climbers 
distributed in Indomalesia and eastern Asia. Although 
this is the first report of fossil Actinidia from North 


34 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


B 


Text-figure 9.—Transverse sections of Alangium endocarps from 
the Nut Beds. A, A. eydei sp. nov., showing relatively thin endocarp 
wall, and one locule somewhat larger than the other. B, A. rotun- 
dicarpum sp. nov., showing thick walls, and locules equal in size. 


America, seeds of the genus occur from the upper Oli- 
gocene to Pliocene of Europe (Kirchheimer, 1957; Do- 
rofeev, 1963; Mai and Walther, 1978; Friis, 1985). 

Actinidia oregonensis seeds are large in comparison 
to the four extant species that I have seen (A. arguta 
Franch. et Sav., A. callosa Lindl., A. chinensis Planch., 
A. strigosa Hook. f. et Thoms.). Whereas the fossils 
average about 4 mm in length, the extant species ex- 
amined, and most of the European fossil specimens, 
are about 2 mm long. Actinidia seeds from the Pliocene 
of the Netherlands, however, range from 1.8 to 4mm 
long (Kirchheimer, 1957). Kirchheimer (1957) placed 
all of the European specimens in a single fossil species, 
A. foveolata Reid and Reid, but Friis (1985) stated that 
the type material of A. foveolata is larger than the Dan- 
ish and Polish material and has a greater number of 
cell rows across the width of the seed. 


Family ALANGIACEAE DC. 


The Alangiaceae are a tropical family with one genus 
and about 20 living species in Africa and from China 
to Australia. Alangium grows as a tree, less often as 
shrubs or lianas. The genus has an excellent fossil rec- 
ord in Europe and North America (Eyde, Bartlett and 
Barghoorn, 1969; Mai, 1970). Although allied to the 
Cornaceae, and placed in the Cornales by Cronquist 
(1981), Alangium differs by the presence of laticifers 
and alkaloids (Eyde, 1988). The genus is divided into 
four sections that are distinguished on the basis of style, 
stigma, stamen, floral vascular system and endocarp 
characters (Eyde, 1968). The paleogeographic history 
of Alangium is reviewed by Kriitzsch (1989). 

The Nut Beds flora includes two species of Alangium 
based upon fruits. Silicified wood of this genus is also 
known from the Nut Beds (Scott and Wheeler, 1982). 


Based upon the presence of scalariform perforation 
plates, small intervascular pits and long vessel ele- 
ments, Scott and Wheeler (1982) determined that 
Alangium oregonensis wood conforms to Metcalfe and 
Chalk's (1950) “group A” (A. javanicum type), and 
differs from “group В” (A. chinense type). This obser- 
vation is of interest because the fruit morphology of 
the species considered below corresponds closely to 
that of extant A. chinense Rehder. 


Genus ALANGIUM Lam. 


Alangium eydei sp. nov. 
Plate 4, figures 5-14, Text-figure 9A 


Etymology.—This species is named for the late 
Richard Eyde, in recognition of his contributions in 
understanding the morphology of living and fossil 
Alangium and other Cornales. 

Description.—Endocarp bilocular, ovate in face view, 
laterally compressed in the plane of septum; base 
rounded, apex obtuse-pointed; length 4.9-10.3, avg. 
7.7 mm (SD=1.7, n=13), width 5.1-7.6, avg. 6.3 mm 
(SD=0.75, n=11), thickness 3.2-6.4, avg. 4.3 mm 
(SD=0.75, n=15); one locule slightly smaller than the 
other; each locule with a single seed; endocarp surface 
relatively smooth with faint broad ridges; position of 
the septum marked on the exterior of the endocarp by 
a peripheral furrow between the two carpels; septum 
ca. 0.4 mm thick, containing six to eight canals that 
radiate in the median plane and terminate before 
reaching the edge of the endocarp, forming an asterisk 
pattern; endocarp wall ca. 0.5 mm thick, made up of 
isodiametric polygonal sclereids 17-30 um; locule lin- 
ing uniseriate, 17.5 um thick, composed of cuboidal 
sclereids. 

Specimens.— Holotype: USNM 424646, endocarp. 
Paratypes: HU 60003, 60004, UCMP 10710, UF 6311, 
6508, 6645, 6646, USNM 354012-354015, 354016 (2 
specimens), 354018, 354019, 354045, 422393, 424645, 
446073, endocarps; UF 9479, USNM 354020, septal 
canal casts; USNM 354017 (20 specimens), isolated 
locule casts. 

Discussion.— Alangium eydei is represented in the 
Nut Beds by occasional endocarp casts (Pl. 4, figs. 5- 
8) and permineralizations (PI. 4, figs. 11—12). The latter 
may be sectioned to reveal internal morphology and 
anatomy (Pl. 4, fig. 13). Frequently all that remains of 
the endocarp is a carbonaceous powder that disinte- 
grates in the field, leaving the two chalcedony locule 
casts side by side, resembling a sandwich (Pl. 4, figs- 
9, 10). Sometimes star-shaped chalcedony casts of the 
canals within the septum are recovered (Pl. 4, fig. 14) 
but the most common remains of Alangium in the Nut 


| 
| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 35 


Beds collections are isolated silica locule casts, pre- 
served as thin, elliptical wafers with a faint stellate 
impression on one side corresponding to the septal 
canals. 

This species, like most of those known from the 
Tertiary (Eyde, Bartlett and Barghoorn, 1969; Mai 
1970), corresponds to section Marlea of extant Alan- 
gium in having a bilocular, thick-walled endocarp. Eyde 
(1968) noted that the extant species of this section can 
be arranged in a sequence to show the gradual loss of 
one ofthe carpels. In Alangium kurzii Craib the locules 
are more or less equal in size, whereas in A. platani- 
Јоћит (8. et Z.) Harms, A. alpinum W. Smith et Cave 
and A. rotundifolium (Hassk.) Bloemb. one of the car- 
ре! is much smaller than the other, often with a tightly 
closed abortive locule, and in A. griffithii, only one 
Carpel is evident. The fossil closely resembles A. chi- 
nense Rehder in having one locule smaller than the 
Other (Text-fig. 9A) , but with both locules fertile. Eyde 
(1968, fig. 3h) illustrated a transverse section of an 
endocarp of extant A. chinense that is almost identical 
to that seen A. eydei (Pl. 4, fig. 13). The smaller of the 
two carpels generally is at least 80% of the size of the 
larger carpel both in the fossil and in A. chinense. 

Alangium eydei bears a close resemblance to A. jen- 
kensii Chandler (1961b) from the London Clay, al- 
though the largest of the Nut Beds specimens is slightly 
Smaller than the single London Clay specimen. Al- 
though A. jenkinsii clearly belongs to Alangium, Chan- 
dler (1961b) noted that “owing to the poor condition 
Of the single specimen it is difficult to define specifi- 
cally." A. vermontanum Eyde et Barghoorn from the 
Oligocene Brandon Lignite of Vermont (Eyde, Bartlett 
and Barghoorn, 1969) differs from А. eydei in having 
a much more reduced smaller locule, and in the retic- 
ulate sculpture of Ше ventral side of the larger locule. 
The Clarno fossil differs from 4. dubium (Unger) Mai 
from the Miocene of central Europe (Mai, 1970, p. 
474) in the lack of transverse ridges on the endocarp 
Surface. 


Alangium rotundicarpum sp. nov. 
Plate 4, figures 15, 16, Text-figure 9B 


Etymology.— Когипаи (L = circular, round), + kar- 
Роз (Gr = fruit), referring to the rounded outline ге- 
Sulting from thick wall of the endocarp. 

Description.—Endocarp subglobose, bilocular; length 
8.5 mm, width 7.2 mm, thickness 7.7 mm; locules 
approximately equal in size, each with a single seed; 
Position of the septum marked at the endocarp surface 
bya peripheral furrow between the two carpels; septum 
0.5 mm thick, containing six to eight canals that radiate 
In the median plane and terminate before reaching the 


edge of the endocarp, forming an asterisk pattern; en- 
docarp wall 2.75 mm thick dorsally and ventrally, thin- 
ner laterally, made up of isodiametric to anticlinally 
elongate sclereids, 40-150 um in diameter; locule lin- 
ing not well preserved, 30 um thick. 

Specimen.— Holotype: USNM 435005. 

Discussion.— This species, known from a single per- 
mineralized specimen, differs from A. eydei in its glo- 
bose endocarp (Pl. 4, fig. 15) with thick lateral walls 
and in the equivalent size of the two locules (Text-fig. 
9B). Although the exterior surface of the endocarp is 
not preserved, the intact portion is about five times 
thicker than the complete wall of A. eydei. Transverse 
sectioning revealed that the locules of A. rotundicar- 
pum are approximately equal in size (Pl. 4, fig. 16), in 
contrast with the unequal development of locules in 
A. eydei (Pl. 4, fig. 13). The endocarp includes anti- 
clinally elongate sclereids, whereas those of A. eydei 
are more or less isodiametric. In size of the endocarp 
and unusually thick endocarp wall, this species resem- 
bles A. krutzschii Mai from the middle Eocene Gei- 
seltal flora of Germany (Mai, 1970); the locules of that 
species, however, are much smaller. 


Family ANACARDIACEAE Lindley 


The Anacardiaceae, or sumac family, includes about 
70 extant genera of tropical to temperate distribution 
including trees, shrubs and lianas, and is divided into 
five tribes: Anacardieae (eight genera), Spondiadeae 
(17 genera) Rhoeae (40 genera), Semecarpeae (five 
genera), Dobineeae (one genus) The Spondiadeae, 
characterized by drupes with hard, often multilocular, 
stones having oval germination apertures, is well rep- 
resented in the fossil record, with records of both extant 
and extinct genera in the early Tertiary of Europe (Reid 
and Chandler, 1933; Collinson, 1983) and North 
America. 

The Anacardiaceae are represented in the Nut Beds 
flora by at least two genera of fruits: Rhus, of the tribe 
Rhoeae, and the extinct genus Pentoperculum of the 
tribe Spondiadeae. In addition, the fruit described 
herein as Pistachioides (p. 105) resembles that of extant 
Pistachia, and might also belong to the Anacardiaceae. 
Anacardiaceous woods from the Nut Beds include 
Tapirira (tribe Spondiadeae; Manchester, 1977), and 
at least one additional genus remaining to be described 
(S. R. Manchester, unpublished). 


Genus PENTOPERCULUM gen. nov. 


Etymology.— Penta (Gr = five) + operculum (L = 
lid), referring to the five germination valves ofthe fruit. 

Type species.—Pentoperculum minimus (Reid et 
Chandler), comb. nov. 


36 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Pentoperculum minimus (Reid et Chandler) 
comb. nov. 
Plate 5 figures 1-17 


Dracontomelon minimus Reid et Chandler 1933, p. 302, pl. 13, figs. 
20-24. 


Description.—Endocarp subglobose to more or less 
oblate, five-(rarely four- or six-) carpellate, roundly 
pentangular (rarely quadrangular or hexagonal) in 
transverse section, length 4.4-6.5, avg. 5.7 mm 
(SD=0.49, n=23), equatorial diameter 4.5-8.5, avg. 
6.8 mm (SD=0.89, n=24), length/equatorial diameter 
ratio 0.74-1.11, avg. 0.90, widest at the equator; base 
round to flat with a slight depression ca. 1 mm in 
diameter at the extreme base, apex rounded; locules 
five (rarely four or six), single-seeded, radially ar- 
ranged, vertically elongate, 3.5-5.0 mm long, 1.5-2.1 
mm wide, 2.1—3.3 mm in dorsiventral dimension; each 
locule with a convex, dorsal-apical, germination valve, 
resulting in five (rarely four or six) elliptical bulges at 
the surface of the endocarp extending from the equator 
almost to the apex; germination valves 440 um thick, 
two-partite with a median longitudinal slit of shutter- 
like splitting; lacunae alternating with the locules at 
the equator of the fruit corresponding in position to 
the exterior angles of the fruit; these lacunae about as 
wide as the locules, but shorter both dorsiventrally and 
axially; central axis of the fruit with a cylinder of vas- 
cular tissue extending from base spreading and dissi- 
pating apically; endocarp wall and septa composed of 
isodiametric sclereids 28-45 um in diameter with in- 
terspersed tracts of fibers; locule lining 25-30 um thick, 
composed of horizontally aligned fibers, these fibers 7 
um in diameter and at least 10 um long. 

Specimens.—UCMP 10633, UF 5708-5711, 6608- 
6610, 10634, UM 31604, USNM 312762 (Bones, 1979, 
pl. 3, fig. 6), 353967 (10 specimens), 353969, 422390, 
424851, 435106-435114, endocarps; UCMP 10635- 
10637, UF 6664-6671, 9317, 9736 (100 specimens), 
USNM 435115 (7 specimens), isolated locule casts. 
Holotype: BM v22546 from the London Clay. 

Discussion. —Specimens of Pentoperculum are pre- 
served in the Nut Beds as white chalcedony casts of 
the endocarp and as permineralized endocarps with 
intact seeds. Most specimens are pentalocular (e.g., Pl. 
5, figs. 1, 2, 8), although two specimens are four-loculed 
(e.g., Pl. 5, fig. 15) and three are six-loculed (USNM 
435107-435109). In some cases the endocarp has dis- 
integrated leaving a cavity in the matrix with protrud- 
ing chalcedony locule casts. Isolated locule casts are 
common (Pl. 5, figs. 5, 6). Because the fossil does not 
correspond precisely to any living genus, the new ge- 
neric name Pentoperculum is established to accom- 
modate this species. Pentoperculum minimus was a 


widespread species that occurred in the Eocene of Hun- 
gary (Kovacs, 1957), as well as England (Reid and 
Chandler, 1933) and western North America. 

This species was first recognized on the basis of pyr- 
itized endocarps and locule casts from the London Clay 
flora (Reid and Chandler, 1933), an example of which 
is included in Pl. 5, fig. 14. The Nut Beds specimens 
are indistinguishable both in external and internal 
morphology from the type material. Reid and Chan- 
dler (1933) demonstrated similarities to the extant ge- 
nus Dracontomelon, and their work confirmed affini- 
ties with the tribe Spondiadeae. However, new 
observations based upon sections of the fossil and ex- 
tant endocarps indicate significant differences between 
Pentoperculum and Dracontomelon. Contrary to the 
interpretation of Reid and Chandler (1933), serial sec- 
tions (e.g., Pl. 5, figs. 9, 10) indicated that the lacunae 
are not linked to peripheral apertures in the fossil, a 
condition considered diagnostic of the extant genus. 
The fossil fruits are more symmetrical than those of 
Dracontomelon, with more or less equal development 
of locules. In addition, the fossil endocarps are much 
smaller than those of the living species of Draconto- 
melon. In contrast to Dracontomelon, in which each 
of the locules opens by the “uncorking” of an undi- 
vided apical plug (Hill, 1933), the locules of Pento- 
perculum open by means of a bipartite operculum (Pl. 
5, fig. 8). 

Various modes of germination occur in the fruits of 
the Spondieae and are described in detail by Hill (1933, 
1937). The apertures of Pentoperculum appear to have 
had shutter-like valves similar to those observed in 
Haematostaphis (Hill, 1933) and Pseudospondias (Hill, 
1937). In these genera, “the lid separates into two equal 
halves which are pushed apart and thrown aside by the 
radicle as it emerges” (Hill, 1933, pp. 881—882). 


Genus RHUS L. 


Rhus rooseae sp. nov. 
Plate 6, figures 1—8 


Etymology.— This species is named after Carrie L. 
Roose, recognizing her helpful assistance and support 
of this project. 

Description.—Fruit bilaterally symmetrical, subel- 
liptic, unilocular, laterally flattened in the plane of 
symmetry, transversely elliptic in cross section, base 
and apex obtuse-rounded; width greater than height; 
height 3.3-4.3, avg. 3.8 mm (SD=0.29, n=8), width 
4.2-5.1, avg. 4.7 mm (SD=0.4, n=5), thickness 1.5- 
2.0, avg. 1.8 mm (SD=0.25, n=9); mesocarp with 25- 
30 prominent longitudinal vascular bundles; endocarp 
surface smooth, shiny, with a circular funicular de- 
pression in the plane of symmetry, and a slight bulge 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 37 


on the opposite side; endocarp wall 400 um thick, com- 
posed of three layers: inner layer 250 um thick, uni- 
Seriate, composed of anticlinally oriented columnar 
cells 250 um high and 20-30 um wide; the next layer 
much thinner, also uniseriate, 33 um thick, composed 
of rectangular cells typically 45 um wide; the overlying 
layer uniseriate, composed of columnar cells similar 
to the first, but only 80 um thick, capped by a uniseriate 
to triseriate layer of rectangular cells. 

Specimens.— Holotype: USNM 355036. Paratypes: 
UF 9350-9352, 9417-9422. 

Discussion.—Sumac fruits, Rhus rooseae, typically 
are preserved in the Nut Beds as translucent, silicified 
endocarps. The prominent columnar cell construction 
is exhibited both in fractured specimens by SEM (PI. 
6, figs. 5, 6) and with transmitted light in thin sections 
(Pl. 6, fig. 8). Only one specimen, designated the ho- 
lotype, has the mesocarp intact showing the prominent 
longitudinal vascular bundles (Pl. 6, figs. 1-3). 

Fruit and seed morphology of Rhus are reviewed by 
von Teichman and Robbertse (1986). R. rooseae shares 
many characters with extant Rhus including lateral 
compression in the plane of bisymmetry, width greater 
than height, longitudinal vascular bundles forming ribs 
in the mesocarp, and endocarp comprised of two or 
more layers of anticlinally elongate columnar cells. Al- 
though Rhus has been reported commonly on the basis 
Of leaves from the North American Tertiary, this is the 
first North American report based upon fruits. The 
genus has also been documented on the basis of fruits 
Similar to extant Rhus toxicodendron L. from the Mid- 
dle Miocene of Denmark (Friis, 1979). The Danish 
fossil differs from R. rooseae in possessing only about 
12, rather than 25-30, longitudinal ribs. 


Family ANNONACEAE Juss. 


The Annonaceae are a family of trees, shrubs and 
lianas with about 128 living genera and 2050 species. 
The family is mostly tropical, with the exception of 
few genera that extend into temperate regions. Three 
Species of Annonaceae can be identified from the Nut 
Beds on the basis of internal molds of the seed coat, 
Showing the characteristic ruminate endosperm. 


Genus ANONASPERMUM Ball 
emend. Reid et Chandler 


Reid and Chandler (1933) emended the fossil genus 
Anonaspermum Ball to accommodate anatropous seeds 
with ruminate endosperm of the various types found 
Ш the family Annonaceae, and with an encircling band 
Of fibers formed by the raphe and chalaza, which di- 
Vides the seed into more or less symmetrical halves, 
With the hilum terminal. Most of the fossils that have 
been attributed to this genus are internal molds of the 


seed coat, or endosperm casts, with the seed coat miss- 
ing. Distinctive surface patterns result from the con- 
figuration of the ruminate endosperm. 

The Clarno specimens of Anonaspermum have been 
segregated into three species using criteria similar to 
those used in distinguishing London Clay species. 
Fourteen species of Anonaspermum were described 
from the London Clay (Reid and Chandler, 1933; 
Chandler, 1961b; summarized, Collinson, 1983), based 
upon differences in seed shape, pattern of ridges in 
endosperm, and nature of the encircling fiber band. As 
noted by Collinson (1983, pp. 73-74), “Many different 
fossil forms may be recognized but it is very difficult 
to relate these to living genera. Similar endosperm pat- 
terns may occur in different living genera and different 
endosperm patterns may occur within the same living 
genus”. It is possible that the three Nut Beds species 
recognized here were actually produced by different 
genera, but because of the limited systematic resolution 
provided by seed characters in this family, they are all 
be placed in the stereotypic genus Anonaspermum. 


Anonaspermum cf. pulchrum Reid et Chandler 
Plate 6, figures 9-15 


Description. —Seed bilaterally symmetrical, short- to 
long-ellipsoid, all faces convex rounded, sometimes 
slightly flattened laterally in the plane of symmetry; 
length 5.5-10.3, avg. 8.0 mm (SD=1.47, n=10), width 
3.8—6.2, avg. 5.2 mm (SD=0.85, n=12), thickness 3.4— 
4.9, avg. 4.3 mm (SD=0.42, n=12); without a marked 
longitudinal median depression; internal mold of seed 
coat with rumination ridges many, narrow, transverse, 
interrupted in course; these ridges mainly continuous 
across the center, without forming pits, nodules or 
punctae; sometimes with short grooves along the ridge 
crests; ruminations tetrapartite in transverse section; 
with a strap-like ridge 1-1.4 mm wide in the plane of 
symmetry formed by fibers of the raphe and chalaza; 
hilum terminal at the broader end of the raphe ridge; 
seed coat thin (80-100 um), smooth. 

Specimens.— HU 60061, UCMP 10676, 10677, UF 
5233, 5234, 6574, USNM 353971 (7 specimens), 
353972, 353975, 353976, 424794-424797, 434983. 

Discussion. — This species is represented in the Nut 
Beds mostly by internal molds of the seed coat (i.e., 
endosperm casts). In one specimen (UCMP 10677) 
part of the thin seed coat is adhering. In seed shape, 
size and pattern of endosperm ruminations, this spe- 
cies corresponds closely to Anonaspermum pulchrum 
Reid et Chandler from the London Clay flora. Reid 
and Chandler (1933) described the raphe and chalaza 
as lying in a rather inconspicuous marginal groove, but 
in most of the Clarno specimens these form a ridge 
(Pl. 6, figs. 10, 13). Perhaps this difference is due to 


38 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


abrasion of the London Clay specimens resulting in 
loss of the fibrous band. In some of the Nut Beds 
specimens, the fibrous band is partially or entirely bro- 
ken away (e.g., Pl. 6, fig. 15) revealing a marginal groove 
like that of the London Clay species. Reid and Chan- 
dler (1933) called attention to similarity with the living 
Indomalaysian species Dasymaschalon clusiflorum 
Merrill, but refrained from suggesting a close relation- 
ship because of the likelihood that similar seeds could 
be found elsewhere in the family with a more exhaus- 
tive search of modern genera. 


Anonaspermum bonesii sp. nov. 
Plate 6, figures 16-18 


Etymology.— This species is named for the late Tho- 
mas J. Bones, who collected most of the Anonasper- 
mum specimens known from the Nut Beds. 

Description.—Seed roughly ellipsoidal, pointed api- 
cally, with a slight flattening near the hilum, rounded 
basally, bilaterally symmetrical, laterally compressed 
in the plane of symmetry such that the thickness is less 
than Y of the width, with some shallow depressions 
on the flattened face; length 11.7 mm, width 7.4 mm, 
thickness 2.5 mm; internal mold of seed coat with 
ruminations transverse in the midsection of seed, but 
radially oriented near the base and apex, thin, closely 
spaced, irregular, disappearing toward the center of the 
face, giving rise to fine punctae; inner layer of seed coat 
with a criss-cross pattern of straight to somewhat curved 
fibers. 

Specimen.— Holotype: USNM 353975. 

Discussion.—This species, known from a single si- 
licified seed, differs from A. cf. pulchrum in its pro- 
nounced lateral compression (Pl. 6, fig. 18), such that 
the thickness is less than % of the width, and by the 
orientation of ruminations. Although the ruminations 
are transversely oriented across most of the seed in A. 
cf. pulchrum (Р1. 6, figs. 9, 10, 14), they are radially 
oriented in the apical and basal 3 of the A. bonesii 
seed (Pl. 6, figs. 15, 17). 


Anonaspermum rotundum sp. nov. 
Ра е Stress 2 


Etymology.— Rotundus (L = circular, round) refer- 
ring to the seed shape. 

Description.—Seed wide-ellipsoidal, rounded api- 
cally and basally, laterally compressed in plane of the 
raphe; length 13.5 mm, width 12.0 mm, thickness 6.0 
mm; internal mold of seed coat with ruminations ra- 
dially oriented from the center, thin, short, irregular, 
shorter toward the center; raphe band extending nearly 
350° around periphery of the seed. 

Specimen.— Holotype: USNM 435099. 

Discussion.—A. rotundatum is known from a single 


specimen which is preserved as an internal mold of 
the seed coat. The specimen differs from those of the 
other two Clarno species by its larger size and nearly 
circular outline. 


Family ARALIACEAE Juss. 


The Araliaceae, as traditionally circumscribed, are 
a family of about 57 genera and 800 species including 
trees, shrubs and lianas mostly of tropical distribution. 
The more cosmopolitan, mostly herbaceous Apiaceae 
(Umbeliferae) with 418 genera and 3100 species, are 
very similar and should be included within Araliaceae 
to have a monophyletic family (W. Judd, pers. comm., 
1991; and see Mabberley, 1989). 


Genus PALEOPANAX gen. nov. 


Etymology.— Paleo (Gr = old) + panax (L, Gr = 
ginseng) referring to the ginseng family, Araliaceae. 
Type species.— Paleopanax oregonensis sp. nov. 


Paleopanax oregonensis sp. nov. 
Plate 7, figures 3-4 


Description.— Fruit wide-elliptical in face view, bi- 
carpellate, bilaterally symmetrical, dorsiventrally 
compressed, with an epigynous perianth bulge; base 
flat or somewhat cordate, apex rounded; length 4.5- 
5.6 mm, width 5.4—6.6 mm, estimated thickness 1—2 
mm; carpels D-shaped in face view, with two to three 
arched longitudinal grooves, adjoined ventrally at the 
central fruit axis by their straight margins, outer mar- 
gins convex, smooth; pedicel > 2.0 mm long, 70.3 mm 
thick; two styles arising parallel to each other from the 
apex, 2.0-3.0 mm long, recurving distally. 

Specimens.— Holotype: OMSI Pb810. Paratypes: 
OMSI Pb1779, Pb1014. 

Discussion.— This species, represented only by com- 
pression fossils, was recovered from the leaf horizon 
at the base of Face 3 in the Nut Beds. The preservation 
of the styles indicates that they were tough and per- 
sistent, or that the fruits were deposited without rig- 
orous transport. 

Fruits of Paleopanax conform to those of the Apiales 
(Araliaceae + Apiaceae) in the possession of epigynous 
perianth, styles as many as the carpels and schizocarp 
morphology. Although samaroid schizocarps also oc- 
cur in Sapindaceae and Aceraceae, the epigynous tepals 
rule out these families. In addition the apical bulge in 
the perianth region (Pl. 7, fig. 3) may be interpreted as 
a nectary disk, as is characteristic of Apiales. Paired 
thin mericarps are characteristic of Apicaceae and also 
occur in some Araliaceae. Specialized carpophores of 
the type found in Apiaceae do not appear to be de- 
veloped in this fossil. 

Similar fruits occur in the extant genus Pseudopan- 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 39 


ax. Fruits of P. davidii (Franch.) W.R. Philipson from 
western Hupeh, China (Pl. 7, fig. 5) are particularly 
similar to the fossil. However, Pseudopanax has a sin- 
gle style that divides distally into two arms, whereas 
the fossil has two distinct styles. 


Family BETULACEAE Gray 


The Betulaceae are a family of six extant genera, 
with an excellent fossil record in the northern hemi- 
sphere (Crane, 1989). The family is represented in the 
Nut Beds by the two fruit types described below, Cor- 
yloides and Kardiasperma, and by wood attributed to 
Betula (Scott and Wheeler, 1982). The family is rep- 
resented at other Clarno Formation localities, includ- 
ing West Branch Creek and White Cliffs, by leaves and 
infructescences of Alnus and by fruits that may be sim- 
ilar to Palaeocarpinus. Also a fruit resembling a small 
Shuttle-cock, Calycites ardtunensis Crane, is known 
from West Branch Creek, White Cliffs and Gosner Road 
localities of the Clarno Formation, and may represent 
an extinct genus of the Betulaceae (Crane, 1989). 


Genus CORYLOIDES gen. nov. 


Etymology.—The generic name refers to the simi- 
larity of these fossils to nuts of Corylus. 
Type species.—Coryloides hancockii sp. nov. 


Coryloides hancockii sp. nov. 
Plate 7, figures 6-12, 16-18 


Etymology.—The species is named in honor of the 
late Alonzo W. Hancock, amateur fossil collector and 
founder of the field station adjacent to the Nut Beds 
locality. 

Description.—Nut nearly perfectly spherical, round- 
€d apically and basally; unilocular, single seeded; out- 
ermost surface as seen in casts and external molds 
Strongly ribbed meridionally; ribs spaced 2—4 mm apart 
at the equator; fruit wall 0.8-1.2 mm thick with a large 
longitudinal vascular bundle corresponding to each of 
the external ribs. Nut cast spherical, length 24.8-37.5, 
avg. 28.8 mm (SD=2.65, n=30), equatorial diameter 
23.0-32.6, avg. 27.5 mm (SD=2.48, n=30), with a 
Prominent basal scar about 20-25 mm in diameter and 
а circular stylar protrusion 1.5-2.5 mm in diameter at 
the apex; surface of nut cast above the basal scar with 
faint longitudinal ridges corresponding in position to 
the ribs on the fruit surface; perimeter of the basal scar 
forming a circumscissle band ca. 2—3 mm wide with a 
Smooth, circular upper margin at which point the lon- 
8ltudinal surface ribs begin, and a more jagged lower 
Margin with the broken ends of vascular bundles cor- 
responding to the surface ribs, where they may have 
been torn away from the involucre; nut surface slightly 


recessed over the basal scar. Seed more or less globose, 
ca. 20-24 mm in diameter, longitudinally wrinkled, 
with a large circular hilum, about 9.5 mm in diameter 
at one end. 

Specimens.—Holotype: UF 8497. Paratypes: UF 
8492-8496, 8499, 9898, USNM 40544 (Scott, 1954), 
354408, 354425, 354620, 354630, 354632, 354634- 
354636, 354641-354643, 354645, 354647, 354650, 
354654, 354658, 354677, 354757, 354811, 354903, 
354922, 354984, 354986, 354992, UWBM 35268, 
35269, nut casts; UF 8498, nut with seed partially 
exposed; UM 29942, seed. 

Discussion. — Coryloides is distinguished by a nearly 
spherical, meridionally ridged nut with a large circular 
circumsissle mark on one end (Pl. 7, figs. 7, 8, 9, 10) 
and a small stylar scar at the other (Pl. 7, figs. 6, 8). 
The chalcedony-replaced nuts are attractive specimens 
with the shape and density of a large game marble. 
The fruits occur mainly at the top of Faces 3 and 4 in 
the Nut Beds, and are not known from other localities 
of the Clarno Formation. Although these specimens 
were informally identified as palm fruits (Gregory, 
1969; Bones, 1979), the large circular band at the basal 
end of the nut is inconsistent with the morphology of 
extant Palmae. The cellular anatomy of the nutshell 
usually is not preserved but in one of the more com- 
pletely preserved, although somewhat crushed, speci- 
mens, transverse sectioning reveals the thickness and 
sculpture of the outer wall, and shows the presence of 
a large vascular bundle corresponding in position to 
each of the surface ribs (Pl. 7, figs. 17, 18). 

Scott (1954) figured a fruit cast (p. 86, pl. 16, fig. 24) 
and an isolated seed cast (p. 87, pl. 16, figs. 29, 30) of 
Coryloides hancockii among his incertae sedae. A si- 
licified fruit that has been fractured, revealing an intact 
seed cast (Pl. 7, fig. 12) provides the basis for linking 
isolated seed casts, such as that figured by Scott, as 
belonging to the same species as the fruit casts. The 
sedimentary mold surrounding the silicified nut cast 
shows that the external surface was more strongly ribbed 
(Pl. 7, figs. 11, 16) than the nut cast alone would sug- 
gest. The large basal scar indicates probable detach- 
ment from an involucre as is seen in the nuts of various 
Fagaceae and Betulaceae. 

The globose form, unilocular, single-seeded struc- 
ture and prominent basal scar of these nuts at first 
suggested affinities with cupulate fruits of Quercus and 
Lithocarpus in the Fagaceae. Fruits with such promi- 
nent longitudinal ridges are not known in the Fagaceae, 
and the peculiar, more or less crenate pattern of vas- 
cular bundles seen on the basal scar is unlike anything 
observed among extant fagaceous fruits (Kaul, pers. 
comm., 1989). In addition, the nutshells of Quercus, 
Lithocarpus and other Fagaceae do not contain large, 


40 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


widely spaced longitudinal vascular bundles as seen in 
this fossil. 

The affinities of this fruit appear to be close to Cor- 
ylus in the Betulaceae. The nature ofthe scar left behind 
on the nut of Corylus, where Ше involucre, or “hull”, 
detaches (Pl. 7, figs. 13, 14), is very similar to that seen 
in Coryloides. This feature, together with the presence 
of a single locule and seed and the relatively thin nut- 
shell containing unusually large longitudinal vascular 
bundles (Pl. 7, fig. 15), is consistent with the fossil nuts. 
However, extant Corylus fruits are smaller and not as 
perfectly globose as those of Coryloides. The basal scar 
of Coryloides is nearly circular, whereas that of Corylus 
usually somewhat distorted and elliptical, indicating a 
difference in the symmetry of involucral attachment. 
These features, and the prominently ribbed surface of 
the nuts indicate that the fossils do not belong in the 
extant genus. Whether Coryloides is a sister genus to 
Corylus, or a case of parallel evolution in another line 
of the Betulaceae is open to speculation. 


Genus KARDIASPERMA gen. nov. 


Etymology.— Kardia (Gr = of the heart) + sperma 
(Gr — seed). 
Type species.— Kardiasperma parvum sp. nov. 


Kardiasperma parvum sp. nov. 
Plate 8, figures 1—5 


Etymology.— Parvum (L — little) referring to the small 
size of the locule casts. 

Description.— Locule casts small, dorsiventrally 
compressed, obcordate, quadrilaterally symmetrical, 
base rounded or faintly cordate with hilum central, 
apex blunt-pointed in face view (ca. 90°); length 1.3— 
2.4, avg. 2.1 mm (SD=0.23,n=19), width 1.0—2.0, avg. 
1.7 mm (SD=0.25, n=19), thickness 0.5-1.2, avg. 1.0 
mm (SD=0.14, n=19); surface smooth with a keel in 
the major plane ofsymmetry and with a median groove 
running dorsally and ventrally from base to apex in 
the minor plane of symmetry. 

Specimens.— Holotype: USNM 435077. Paratypes: 
UCMP 10710-10713, UF 6602-6604, USNM 355706- 
355710, 424705-424710, 424750, 424768, 435076, 
435078—435081. 

Discussion.— These specimens are morphologically 
identical to the locule casts of Palaeocarpinus from the 
Paleocene of Almont, North Dakota (Crane et al., 1990, 
p. 24). There is a marked difference in size, however. 
Locule casts from the Almont locality are 45.5 mm 
wide, i.e., two to five times larger than the species 
described above. Although the locule casts of Kar- 
diasperma parvum have not been found intact within 
complete fruits, they correspond in size to fruits of an 
undescribed species of small, spiny-bracted fruit sim- 


ilar to Palaeocarpinus known from compressions (Pl. 
8, fig. 20) at the West Branch Creek and White Cliffs 
localities of the Clarno Formation. Palaeocarpinus is 
an extinct genus of the Coryleae known from the Pa- 
leocene of England (Crane, 1981) and North America 
(Crane et al., 1990; Sun and Stockey, 1992). 


Family BURSERACEAE Kunth. 


The Burseraceae, or torchwood family, comprises 
16 genera and about 500 species of tropical trees and 
shrubs with present centers of diversity in America 
and northeastern Africa. The family is divided into 
three tribes (Lam, 1931, 1932; Daly, 1989), based prin- 
cipally upon characters of the fruits. In the tribe Bur- 
sereae (Bursera, Commiphora, Aucoumea, Boswellia) 
pyrenes are concrescent but not connate, and have a 
dry, dehiscent pericarp. In the Protieae (Protium, Ga- 
ruga, Tetragastris, Hemicrepidospermum, Crepidos- 
permum) pyrenes are separate and the outer pericarp 
may be either fleshy or dry and dehiscent. In the Can- 
arieae (Canarium, Scutinanthe, Dacryodes, Santiria, 
Trattinickia, Haplobus) pyrenes are connate, forming 
a plurilocular endocarp with a fleshy, rarely more or 
less dry, indehiscent pericarp. General fruit and seed 
morphological characters for the family are reviewed 
by Reid and Chandler (1933, pp. 266-7). 


Genus BURSERICARPUM Reid et Chandler 


Bursericarpum oregonense sp. nov. 
Plate 8, figures 6-17 


Etymology.— The epithet refers to Oregon, the state 
in which the Clarno fossils were collected. 

Description.— Endocarp single-seeded, bilaterally 
symmetrical, tear-dropshaped, pointed apically, 
rounded basally, subtriangular in cross section, with a 
convex, rounded dorsal face and two more or less flat 
ventral faces meeting in a median longitudinal angle 
of 70—90°, this angle sharper above the placenta; length 
4.7-8.7, avg. 6.0 mm (SD=0.97, n=42), width 2.8—5.7, 
avg. 4.3 mm (SD=0.82, n—42), thickness 2.2-4.9, avg. 
3.7 mm (SD=0.85, n=42); transverse placental slit 
concave upwards, located centrally on the ventral angle 
of the endocarp; endocarp wall 0.3 mm thick, com- 
posed of more or less isodiametric sclereids 15-35 um 
in diameter; exocarp not anatomically preserved, but 
showing a reticulum of coarse vascular bundles de- 
veloped on dorsal and ventral surfaces; the bundles of 
this reticulum becoming closer together and forming 
a complex at the placental area. Locule casts similar 
in shape to the endocarps, with more pronounced ven- 
tral concavity in the position of placental slit, a sharper 
(less rounded) ventral angle, a more pointed apex, and 
a curved ridge near the base of the dorsal side corre- 
sponding to the edge of the germination valve. 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 41 


Specimens.—Holotype: USNM 424885. Paratypes: 
HU 60010, UCMP 10655-10660, UF 6647-6648, 
6657-6663, USNM 326717 (Bones, 1979, pl. 3, fig. 7, 
lower left), 326718 (Bones, 1979, pl. 3, fig. 7, lower 
right), 355491, 424877-424884, 424886-424888, 
434951-434960, 434964. 

Discussion.— Bursericarpum oregonense is герге- 
sented by numerous chalcedony locule casts (Pl. 8, figs. 
16, 17), occasional permineralized endocarps (Pl. 8, 
figs. 6-10, 12, 15), and a few complete fruit casts. The 
rounded angles of the endocarps suggest that they were 
borne freely (not fused) within the fruit. In addition, 
complete fruit casts with the reticulate-veined exocarp 
intact (Pl. 8, figs. 13, 14) show only a single endocarp 
рег fruit. The absence of fruit casts bearing two or more 
endocarps suggests that fruits of this species were typ- 
ically single-seeded, a feature of the tribe Bursereae. 

Among extant genera, Bursericarpum oregonense 
most closely resembles Protium, Tetragastris and Bur- 
Sera. Endocarps of Protium and Tetragastris, of the 
tribe Protieae, generally are larger than the fossils, and 
are often borne two or more per fruit. Bursera endo- 
Carps are borne one-per-fruit and are similar in size to 
the fossil. This species is particularly similar in en- 
docarp morphology to the London Clay fossil Burser- 
icarpum aldwickense Chandler (1961b). A separate 
Species name is necessary, however, because exocarp 
vasculature, and the number of pyrenes per fruit, are 
unknown in B. aldwickense and because features of the 
Seed, readily apparent in the London Clay material, 
are not known for the Nut Beds specimens. 


Bursericarpum sp. 
Plate 8, figures 18, 19 


Description.—Endocarp single-seeded, bilaterally 
Symmetrical, tear-drop-shaped, pointed apically, 
Tounded basally, subtriangular in cross section, with a 
Convex, rounded dorsal face and two more or less flat 
ventral faces meeting in a median longitudinal angle 
of 70—90°, length 2.0-3.2 mm, avg. 2.8 mm (SD=0.5, 
1—5), width 1.3-2.3 mm, avg. 1.9 mm (SD=0.4, n=5), 
thickness 1.0-2.0, avg. 1.6 mm (SD=0.4, n=5); pla- 
Cental slit transverse, located centrally on the ventral 
angle of the endocarp, curved apically through the en- 
docarp. 

Specimens.—UF 6654-6656, USNM 434961, 
434963. 

Discussion. — This species is based upon specimens 
that are morphologically very similar to Bursericarpum 
Огевопепзе, but which are much smaller. Most are pre- 
Served as chalcedony casts of the endocarp; locule casts 
have not been observed. It is possible that these spec- 
Imens may be immature or abortive endocarps of B. 
Oregonense. 


Family CORNACEAE Dumort. 


The Cornaceae, or dogwood family includes six ex- 
tant genera: Cornus, Mastixia, Nyssa, Camptotheca, 
Davidia and Diplopanax (Eyde, 1988; Eyde and Xiang, 
1990). Although Nyssa and Davidia have often been 
segregated as Nyssaceae, Eyde (1988) reviewed the cri- 
teria used to separate Cornaceae and Nyssaceae and 
found them inadequate to support the continued rec- 
ognition of Nyssaceae. Most species of the Cornaceae 
are arborescent and are distributed from temperate to 
tropical regions. The Nut Beds assemblage includes 
both extant and extinct genera of the family: Cornus, 
Mastixia, Mastixioidiocarpum, Mastixicarpum, Lang- 
tonia and Nyssa. Other relatives of Cornaceae present 
among the Nut Beds fruits include Alangiaceae (p. 34) 
and Hydrangeaceae (p. 50). 


Genus CORNUSL. 


Cornus clarnensis sp. nov. 
Plate 9, figures 1-15 


Etymology.— The epithet is named after Clarno, Or- 
egon, the small settlement west of the Nut Beds. 

Description. — Endocarp nearly globose to ellipsoid- 
al, bilocular (to trilocular), length 4.2, 4.4, 5.0 mm, 
width 3.8, 4.4, 4.3 mm; length/width ratio 1.0-1.3; 
base rounded, apex obtusely pointed; surface with ca. 
10 fine longitudinal grooves; endocarp wall 0.5-0.6 
mm thick, without lacunae, composed of more or less 
isodiametric polygonal sclereids, 38-70 um in diam- 
eter, decreasing in size toward the outer surface; sep- 
tum 0.2-0.25 mm thick without central vascular strand 
but with two peripheral vascular strands forming 
prominent longitudinal grooves along the endocarp 
surface in plane ofthe septum; locule lining uniseriate, 
composed of thin-walled cuboidal cells 45 um in di- 
ameter along the distal surface ofthe locule and 25 um 
on the proximal surface (nearest the septum); each loc- 
ule with one seed. 

Specimens.— Holotype: USNM 422378. Paratypes: 
UF 6310, 9763, USNM 355397, 424702. 

Discussion.— Cornus clarnensis is represented only 
by five specimens: a chalcedony endocarp cast and four 
permineralized endocarps. Among the four specimens 
with internal structure preserved, three are bilocular 
(e.g., Pl. 9, figs. 7, 8) and the other is trilocular, but 
otherwise identical in anatomy (Pl. 9, fig. 12). The lack 
of a central vascular bundle in the septum supports 
placement in Cornaceae. The species corresponds to 
extant Cornus in the smooth ellipsoidal outline of the 
endocarp, the number of locules and the isodiametric 
sclereids making up the endocarp. Based upon analysis 
of many characters of extant species, Eyde (1988) di- 
vided Cornus into two groups that he named infor- 


42 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


B 


D 
С 


Text-figure 10.— Mastixioid fruits from the Nut Beds. A, Mastixia sp. В, Mastixioidiocarpum oregonense Scott. С, Mastixicarpum occidentale 
sp. nov. D, Langtonia bisulcata Reid and Chandler. Epicarp stippled where preserved. Scale bar = | cm. 


mally according to the color of their fruits, the Blue 
line (ca. 50 spp.), and Red line (ca. 15 spp.). In shape 
and distribution of surface grooves, C. clarnensis re- 
sembles extant C. florida L., a member of the “big- 
bracted” subgroup of the “showy bract” group of the 
red line dogwoods (sensu Eyde, 1988). The fossil is 
distinguished from endocarps of the Cornelian cherry 
subgroup (or “dunstania” type) by the absence of la- 
cunae in the endocarp wall. 

A thorough review ofthe fossil record of Cornus was 
presented by Eyde (1988). Cornus clarnensis appears 
to be the oldest known fruit record of the big-bracted 
dogwoods. However, multilocular Cornus endocarps 
of the dunstania type occur in the late Paleocene of 
North Dakota (Crane et al., 1990) and in the early 
Eocene London Clay flora (Eyde, 1988). Although Reid 
and Chandler (1933) attributed the London Clay spe- 
cies to an extinct genus, Dunstania, Eyde (1988) argued 
that the *dunstanias" belong within Cornus and, be- 
cause of lacunae in the endocarp wall, correspond to 
the Cornelian cherry subgroup. 


Genus MASTIXIA Blume 


Mastixia is an extant tropical genus of 13 species 
distributed in Indomalesia. Several fossil species of 
Mastixia (reviewed by Kirchheimer, 1957; Mai, 1970) 
and closely related extinct genera (Kirchheimer, 1936, 
1957) are common in the middle to late Tertiary of 
central Europe, prompting application of the term 
“Mastixoid flora" to these warm-climate fossil assem- 
blages (Kirchheimer, 1940). Knobloch and Mai (1986) 
reported several species of this complex from the Late 
Cretaceous (Maastrichtian) of Germany and presented 
a key distinguishing the European fossil mastixioid 
genera. The mastixioid group is represented in the Nut 
Beds flora by Mastixia, Mastixioidiocarpum, Mastix- 
icarpum, and Langtonia. 


Mastixia sp. 
Plate 10, figures 1-3; Text-figure 10А 


Description.—Endocarp ovoid, unilocular, single- 
seeded, surface smooth; length 8.0-11.5 mm (esti- 
mated from incomplete specimens), width 5.7—7.0, avg. 
6.3 mm (SD=5.7, n=5), dorsiventral thickness 6.0- 
6.7, avg. 6.4 mm (SD=0.33, n=4); dorsal surface with 
a sulcus formed by a deep longitudinal infold of the 
endocarp wall extending about 2.8 mm into the locule; 
infold thick (1.8-2.4 mm); locule U-shaped in cross 
section; endocarp wall 0.6-0.8 mm thick, composed 
of two layers: the inner layer 0.3-0.4 mm thick with 
fibers 14-20 um in diameter and more than 400 um 
long, mostly oriented transversely; outer layer approx- 
imately the same thickness, 0.3-0.4 mm, composed of 
more or less isodiametric to elongate sclereids, 20-35 
um wide and up to 200 um long; germination valve 
elongate; dehiscence plane running nearly the full length 
of the endocarp, extending from the dorsal limbs of 
the locule through the thickness of the endocarp wall; 
two longitudinal lacunar tubes up to 0.5 mm in di- 
ameter situated parallel and adjacent to the dehiscence 
plane, one on each side of the locule; epicarp not pre- 
served. Locule cast ellipsoidal, concave dorsally, trough- 
shaped, base rounded, apex pointed; length 7.3 mm, 
width 4.8 mm, thickness 4.1 mm. 

Specimens.—UF 6303, 6502, 9369, 9470, permi- 
neralized endocarps; USNM 354035, locule cast. 

Discussion.— This species, represented by a few per- 
mineralized endocarps and a locule cast, conforms 10 
extant Mastixia on the basis of its relatively smooth 
endocarp surface, the dorsal infold and germination 
valve and the anatomical structure of the епдосагр 
consisting of an inner layer of transversely aligned fi- 
bers surrounded by an outer layer of sclereids (Pl. 10, 
fig. 3). Kirchheimer (1936, pl. 5, fig. 10) and Eyde 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 43 


(1963) illustrated endocarps of comparable living spe- 
cies including M. trichotoma Blume. Eyde (1963) ob- 
served that: “the endocarp splits exactly where two 
Ovular traces pass through the endocarp from base to 
Placenta. These vascular strands form hollow space 
where the ovular trace deteriorates”. The same struc- 
tures are observed in the fossil (Pl. 10, fig. 2; Text-fig. 
10A) and are described above as lacunar tubes. The 
dorsal infold is much thicker in this fossil than in most 
extant species. However, the two species of subgenus 
Manglesia (M. euonymoides Prain and M. octandra 
Matthew), which I have not seen, are described as hav- 
ing thick infolds (“septum swollen to one third the 
diameter of the fruit", Matthew, 1976). 

This species differs from the other Clarno mastixioid 
Species by its smaller size, thinner endocarp wall and 
Smooth surface. The lack of preserved epicarp (me- 
Socarp and exocarp) may indicate that the outer layers 
were fleshy and hence lost prior to fossilization. The 
Smooth surface of the endocarp and small size differ- 
entiates this species from the nodular surfaced endo- 
carp of Mastixia cantiensis and from the gently ribbed 
M. parva from the London Clay flora (Reid and Chan- 
dler, 1933). 


Genus MASTIXIOIDIOCARPUM Scott 


Mastixioidiocarpum oregonense Scott 
Plate 10, figures 4-11, Text-figure 10B 


Mastixioidiocarpum oregonense Scott 1954, р. 84, pl. 16, figs. 16— 
18. 


Emended Description.—Fruit ovoid, rounded api- 
cally and basally, unilocular, single-seeded, length 13- 
23.8, avg. 20.2 mm (SD=3.91, n=6), width 9.2-15, 
avg. 11.7 mm (SD=2.01, n=8), dorsiventral thickness 
8.2-15.0, avg. 11.6 mm (SD=2.45, n=7); endocarp 
longitudinally ribbed with nine to 14 prominent irreg- 
ular sharp ridges up to 1.5 mm in relief that are irreg- 
ularly divided into excrescences or protuberances; en- 
docarp surface with a dorsal sulcus (longitudinal infold) 
€xtending 3.8-5.8 mm into the locule (hence, locule 
U-shaped in cross section); endocarp wall 2.5-4 mm 
thick measured to the crest of the highest ribs; ger- 
Mination valve elongate; dehiscence plane running 
Nearly the full length of the endocarp, extending from 
the dorsal limbs of the locule through the thickness of 
the endocarp wall; two longitudinal lacunar tubes up 
to 0.5 mm in diameter running through the endocarp 
adjacent to the dehiscence plane, one on each side of 
the locule; endocarp consisting of an inner layer 0.3 to 
9.4 mm thick made up of horizontal, periclinally ori- 
ented fibers, 17-23 um in diameter, grading into the 
Outer layer 2.2-3.7 mm thick composed of fibers, 28— 

0 um in diameter, arranged in tracts of varying ori- 


entation; locule lining formed by transversely aligned 
cells; epicarp not preserved. Locule cast narrowly sad- 
Че shaped, the ventral face convex, the dorsal face 
concave, U-shaped in transverse section, the arms 
proximate, lateral margins rounded, but thinning to- 
ward the ends and acute at base and apex, traces of a 
median raphe strand visible on the ventral surface. 

Specimens.— Holotype: UM 29938. Hypotypes: HU 
60013, OMSI Pb196, UF 9206-9208, 9210, 9269, 
USNM 354022-354028 (these specimens together in 
the same piece of matrix), 354029, 354031, 354033, 
354034, 354036, 354037, 354598, endocarps; UCMP 
10661, 10662, USNM 354036, 354038, 354039 (3 
specimens), 354041 (3 specimens) 354042, 354049, 
locule casts. 

Discussion. — Mastixioidiocarpum oregonense is dis- 
tinguished from the other mastixioids of the Nut Beds 
by its prominently sculptured endocarp surface (Pl. 10, 
figs. 4, 8; Text-fig. 10B). It is preserved as perminer- 
alized endocarps, locule casts and endocarp molds. 
Commonly, the endocarp itself has disintegrated, leav- 
ing behind its mold in the matrix, with a chalcedony 
locule cast inside. In one such example, the locule cast 
was removed (Pl. 10, figs. 9—11) and a replica of the 
original endocarp, complete with the irregular surface 
ribs, was prepared by making a latex cast of the sur- 
rounding sedimentary mold (Pl. 10, fig. 8). Molds with 
the same distinctive type of sculpture are known from 
other localities of the Clarno Formation, including West 
Branch Creek, Gosner Road, and from the Eocene Sep- 
ulcher Formation of Yellowstone National Park 
(Princeton 22488, specimen at USNM). 

These endocarps are morphologically similar to those 
of extant Mastixia, but differ by the prominent, coarse 
sculpture and thick fruit wall. Scott (1954) established 
the genus Mastixioidiocarpum for this Nut Beds spe- 
cies, because not enough details were preserved to en- 
able a secure assignment to any of the extinct genera 
known from the European Tertiary (Kirchheimer, 1936; 
Reid and Chandler, 1933). Since Scott's work was pub- 
lished, however, many additional specimens have been 
recovered, some with excellent anatomical detail, en- 
abling a more informed comparison with other fossil 
and modern material. The resulting new information, 
included in the above description, supports retention 
of the generic name Mastixioidiocarpum, because the 
Clarno fossil 1s readily distinguished in its morphology 
from the European fossil mastixioid genera reviewed 
by Holy (1975) and Knobloch and Mai (1986). In 
sculpture and thickness of the endocarp, Mastixioidi- 
ocarpum resembles Eomastixia Chandler (= Ganitro- 
cera Kirchheimer) as emended by Holy (1975). How- 
ever, Eomastixia generally is two-loculed (up to four- 
loculed) and, when unilocular, there is usually evidence 


44 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


ofan abortive ovule (Holy, 1975). Mastixioidiocarpum 
specimens are exclusively unilocular, without any in- 
dication of an abortive ovule. 


Genus MASTIXICARPUM Chandler 


Mastixicarpum occidentale sp. nov. 
Plate 10, figures 12-15; Text-figure 10C 


Etymology.— Occidens (L = of the west), referring 
to the western occurrence of this species. 

Description.—Fruit ovoid, unilocular, length 21.0- 
22.0 mm, width 14.5-16.0 mm, dorsiventral thickness 
11.8-15.5 mm; endocarp with faint longitudinal ridg- 
es; dorsal infold present but not well exposed surfi- 
cially; infold extending 6 mm into the locule; endocarp 
wall 3.6-4.0 mm thick, composed ofinterwoven tracts 
of fibers, the fibers 38-80 um in diameter; vascular 
bundles located in a narrow zone transitional between 
endocarp and epicarp; epicarp surface smooth, exocarp 
0.8-1.0 mm thick, composed of anticlinally elongate 
sclereids, 28—55 um in diameter, 80-300 um long. Loc- 
ule boat-shaped, the dorsal face concave, and the ven- 
tral convex, c-shaped in cross section; dehiscence plane 
running nearly the full length of the endocarp, extend- 
ing from the dorsal limbs of the locule through the 
thickness of the endocarp wall. 

Specimens.— Holotype: UF 8740. Paratype: UCMP 
10705. 

Discussion.—This species is known from two frag- 
mentary but well preserved permineralized specimens 
from the Nut Beds, and by a well preserved specimen 
from the Brummers’s Spring Clarno locality (UF 9886; 
loc. 254). Mastixicarpum occidentale differs from the 
other Clarno mastixoids by the presence of a resistent 
epicarp (Pl. 10, figs. 13-15, Text-fig. 10C). The epicarp 
in the other species was probably fleshy and easily 
degraded as in extant Mastixia. The endocarp of this 
species is smooth in comparison to the strongly ribbed 
endocarps of Mastixioidiocarpum discussed above. 

Among fossil mastixioids recognized in the Tertiary 
of Europe, four genera have persistent epicarps (Knob- 
loch and Mai, 1986): Mastixicarpum Chandler, Mas- 
tixiopsis Kirchheimer, Retinomastixia Kirchheimer, 
and Tectocarya Kirchheimer. Of these, the Nut Beds 
species most closely resembles Mastixicarpum and 
Tectocarya. Tectocarya has a pair oflacunae separated 
by a median septum in the upper % of the dorsal infold, 
and has a thin epicarp that may become free from the 
endocarp, whereas Mastixicarpum lacks the longitu- 
dinal septum in the infold and has a thick epicarp that 
is tightly attached to the endocarp. Based upon sections 
of Tectocarya rhenana Kirchheimer that I have ex- 
amined from Diiren, Germany, there are no prominent 
secretory ducts, and the epicarp is composed of very 


small isodiametric cells. The Clarno species corre- 
sponds most closely in morphology and anatomy to 
Mastixicarpum, based upon sections of M. limnophil- 
um that I have examined from the Miocene of Wiesa, 
Germany. The two species correspond in thickness of 
the epicarp, size and orientation of epicarp cells, and 
in the presence of secretory canals at the junction of 
the endocarp and epicarp. 

Recent recognition of the extant Chinese genus Di- 
plopanax as a living mastixioid (Eyde and Xiang, 1990) 
is of considerable interest in interpreting paleobotan- 
ical remains. Diplopanax, like Mastixia, has an ovoid 
unilocular fibrous endocarp with an elongate dorsal 
germination valve. Although there is a dorsal infold 
resulting in a locule that is u-shaped in cross section, 
the infold is not apparent at the surface of the endocarp, 
which has a circular cross section. Mastixia fruits have 
a soft epicarp, whereas that of Diplopanax is woody 
or leathery. Also, the endocarp of Diplopanax is rid- 
dled with longitudinal secretory canals that are not 
observed in the endocarp of Mastixia. Eyde and Xiang 
(1990) suggested that Mastixicarpum and Diplopanax 
are equivalent. However, in both Mastixicarpum lim- 
nophilum and M. occidentalis the secretory canals are 
mostly peripheral to the endocarp (e.g., Pl. 10, fig. 15), 
largely confined to the epicarp, whereas in Diplopanax 
stachyanthus Hand.-Mazz. the secretory canals are 
uniformly distributed, giving a speckled appearance to 
the endocarp in cross section. 


Genus LANGTONIA Reid et Chandler 


Langtonia bisulcata Reid et Chandler 
Plate 11, figures 1-10; Text-figure 10D 


Langtonia bisulcata Reid et Chandler 1933, p. 453, pl. 25, fig. 18- 
ER 


Description.— Fruit oblong, more or less circular in 
cross section, rounded basally and apically, length 16.0- 
35.0, avg. 24.9 mm (SD=5.27, n=7), width 10.0-26.5, 
avg. 14.8 mm (SD=4.56, п=7), dorsiventral thickness 
9.9-22.0, avg. 14.1, mm (SD=4.12, n=7); bicarpellate, 
with two single-seeded elongate locules, sometimes with 
one of the locules poorly formed, abortive; locules 
w-shaped in cross section, due to a pair of longitudinal 
infolds of the endocarp wall; longitudinal planes of 
dehiscence in the endocarp wall defining a pair of ger- 
mination valves, one corresponding to each locule; each 
germination valve extending the full length of the loc- 
ule, similar in width to the locule that it serves; septum 
without a central vascular column; endocarp made up 
of tracts of fibers oriented in various directions; epicarp 
about 1 mm thick, composed of thin-walled, periclin- 
ally elongate cells 20-30 um high and 40-120 um wide; 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 45 


Secretory ducts present both in endocarp and epicarp; 
seeds conforming in shape to the locule. Locule casts 
elongate, dorsiventrally flattened 12.5-20.6 mm long, 
5.5-12.7 mm wide, 2.2-6.0 mm thick, pointed apically 
and basally, sometimes with fine transverse striations; 
dorsal side with a median longitudinal ridge flanked 
on either side by a prominent groove, ventral side 
convex-rounded with a thread-like median longitudi- 
nal raphe groove or ridge. 

Specimens.—HU 60016, UF 8729-8735, 8740, 
8742, USNM 312746 (Bones 1979, pl. 4, fig. 5 top), 
354032, 354046, 354052, 354053, 354351, 354423, 
354937, 354941, 422524, 424810, 424875, endocarps; 
UF 8736-8739, USNM 312747 (Bones 1979, pl. 4, fig. 
5, left), 354043, 354047, 354050, locule casts. Holo- 
type: BM v22984 from the London Clay. 

Discussion.—Langtonia is represented in the Nut 
Beds by permineralized endocarps (Pl. 11, figs. 1-3) 
and occasional isolated locule casts (Pl. 11, figs. 5, 6). 
Fruits have been recovered from the basal leaf layer 
of Face 3, as well as higher in the Nut Beds section. 
The preservation of fruit morphology and anatomy is 

i et as seen in transverse sections (Pl. 11, figs. 4, 
-10). 

This species was first described and recognized as 
an extinct genus from the London Clay flora by Reid 
and Chandler (1933). They attributed Langtonia to Ше 
Cornaceae and placed it near Mastixia because of its 
long, infolded germination valves. It differs from Mas- 
tixia in having two infolds of the locule, rather than 
One, resulting in a w-shaped, rather than u-shaped, 
Cross section. Kirchheimer (1936) expressed doubt as 
to the mastixioid affinities of this genus. Anatomy re- 
vealed in peels of the well preserved Clarno material 
(Pl. 11, figs. 9, 10), however, shows variously oriented 
tracts of fibers forming the endocarp in conformity 
With the anatomy of Mastixia, Diplopanax and Nyssa. 
In addition, the presence of secretory canals in the 
€ndocarp and epicarp is a feature shared with Diplo- 
Panax and Mastixicarpum. Therefore, Langtonia 
Should be reinstated as a member of the Cornaceae. 

Most of the Nut Beds endocarps are larger than those 
described by Reid and Chandler (1933) from the Lon- 
don Clay (length 9-20, avg. 15.5 mm; width 4.5-12, 
avg. 9.6 mm). The size ranges overlap, however, and 
Ше specimens are morphologically indistinguishable, 
and I have therefore placed the Nut Beds specimens 
Ш the same species. 


Genus NYSSA L. 


Nyssa spatulata (Scott) comb. nov. 
Plate 11, figures 11—19 


Palaeonyssa spatulata Scott 1954, p. 83, pl. 16, fig. 8-12. 


Description.—Endocarp oblong, ovoid or ellipsoid- 
al, trilocular, roughly circular to rounded-triangular in 
cross section, base and apex rounded, with nine round- 
ed longitudinal ridges extending from base to apex, 
length 20.1-30.0, avg. 24.0 mm (SD=3.8; n=15), width 
12.0-21.5, avg. 14.6 mm (SD=2.8, n=16), endocarps 
trilocular, with large, rounded germination valves vis- 
ible as slits in the apical half ofthe endocarp; wall and 
septa composed of swirling tracts of fibers, the fibers 
15-30 um in diameter; central axis not vascularized, 
epicarp not preserved. Locules single-seeded, elongate, 
spatulate, c-shaped in cross section, concave outward, 
broadening and rounded apically, narrowed and point- 
ed basally. 

Specimens.— Holotype: UM 29936 (Scott, 1954, pl. 
16, fig. 8, 9). Paratype: USNM 40543 (Scott, 1954, pl. 
16, fig. 10-12). Hypotypes: OMSI Pb197A, Pb914, UF 
6834-6842, USNM 354541, 354582-354584, 354586, 
354591, 354594, 354595, 354604, 424874. 

Discussion.—This species was described by Scott 
(1954) on the basis of four specimens. Although it is 
not abundant in the assemblage, it is now known from 
approximately 25 specimens, resulting in the broader 
range of dimensions indicated in the above description. 
All of the specimens show three locules (rather than 
three to four as in the London Clay species). 

Reid and Chandler (1933) correctly deduced the close 
affinity of this taxon to extant Nyssa, but because of 
some apparent differences, particularly the presence of 
three to four locules, placed the London Clay fossils 
in a new genus, Palaeonyssa. Scott (1954) recognized 
close similarity between the Clarno and London Clay 
fruits, and placed the Nut Beds specimens in a new 
species of Palaeonyssa. These fossils differ from most 
modern species of Nyssa in being consistently trilocular 
(Pl. 11, figs. 16-18), rather than unilocular, and by their 
larger size. Thin sections reveal that the endocarp is 
made up of swirling fibers (Pl. 11, fig. 19), as is also 
true of extant species of Nyssa. 

Nyssa has about six extant species (three in North 
America, one in China, one in Indomalesia and one 
in southern Central America) that are readily distin- 
guished from one another on the basis of fruit mor- 
phology (Eyde, 1963). The new discovery of an extant 
species of Nyssa from Costa Rica and Panama, N. 
talamancana Hammel and Zamora (1990), is helpful 
in the interpretation of fossil remains. Fruits of N. 
talamancana measure 4—5 cm in length and 2-2.5 cm 
in diameter, and are larger than those of any other 
living or fossil species of the genus. In contrast to the 
other extant species of Nyssa, which typically have only 
one locule, and in rare instances two locules (Eyde, 
1963), stones of N. talamancana are bilocular to tri- 
locular (Hammel and Zamora, 1990). Taking into con- 


sideration the wider range of endocarp morphology 
imparted to Nyssa by the tropical American species, 
the taxon Palaeonyssa now appears to fall neatly within 
the extant genus. 


Nyssa scottii sp. nov. 
Plate 12, figures 1-2 


Etymology.—This species is named for Richard A. 
Scott in recognition of his careful work on the Clarno 
fruit and seed flora. 

Description.—Endocarp spindle shaped, fusiform in 
lateral view, roughly circular in cross section, triloc- 
ular, base and apex bluntly pointed, surface with coarse 
meridional ribs; length 12.6-16.5, avg. 13.8 mm 
(SD=2.3, n=6), width 6.1—9.5, avg. 7.5 mm (SD=1.18, 
n=6), endocarp with three inverted u-shaped germi- 
nation valves over the apical Y of the endocarp; en- 
docarp wall 1.2 mm thick composed of fibers 12-30 
um in diameter, arranged in swirling groups, central 
axis not vascularized. 

Specimens.— Holotype: USNM 354588. Paratypes: 
OMSI Pb207, UF 6843, USNM 354582, 354600, 
435150. 

Discussion. — Nyssa scottii is represented by several 
translucent silica casts and one permineralized speci- 
men (UF 6843). The shape, size, surface sculpture and 
type of germination valve are diagnostic of Nyssa. The 
specimens are tricarpellate as in N. spatulata, rather 
than unicarpellate as is the usual condition in extant 
species. One of the specimens (Pl. 12, figs. 1, 2) has 
one of the germination valves broken away, showing 
a locule that is D-shaped in cross section with the 
convex surface inward. The short length of the ger- 
mination valves is a character that distinguishes Nyssa 
from the mastixioid genera (Chandler, 1926, pp. 36— 
37). This species is readily distinguished from М. spa- 
tulata by its smaller size range (length 12-17, vs. 20— 
30 mm). 


Nyssa sp. 
Plate 12, figures 3—5 


Description.— Fruit unilocular, endocarp more or less 
elliptical in face view, dorsiventrally compressed, 
rounded basally, dorsal side convex, ventral side con- 
cave; length 24.6, 30.0, 48.0 mm (est. from incomplete 
specimen), width 12.9, 23.5, 25.0 mm, dorsiventral 
thickness 5.1, 12.6, 10 mm, surface relatively smooth, 
with thin, shallow longitudinal grooves; endocarp wall 
3.2 mm thick, composed of tracts of swirling fibers, 
the fibers 15-30 um in diameter. Locule cast dorsi- 
ventrally compressed. 

Specimens. — UF 9757, UM 66135, USNM 446088. 

Discussion.— Three unilocular specimens of Nyssa 
have been recovered from the Nut Beds. The longi- 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


tudinal grooves on the endocarp (Pl. 12, fig. 3) and 
thick wall composed of swirling tracts of fibers (Pl. 12, 
fig. 5), are consistent with assignment to Nyssa. There 
is no indication of abortive locules and, in the absence 
of any bilocular specimens in the Nut Beds collections, 
it seems likely that this represents a species distinct 
from the trilocular species, N. spatulata and N. scottii. 
Although it is incomplete, the larger of the specimens 
(Pl. 12, fig. 3) appears to have been longer than any of 
the trilocular specimens. 

The relatively smooth surface, size, and marked dor- 
siventral compression of these endocarps suggests af- 
finities with N. brandoniana (Lesquereux) Eyde and 
Barghoorn, from the Brandon lignite of Vermont (Eyde 
and Barghoorn, 1963) and extant N. javanica (Bl.) 
Wangerin of southeast Asia and Malesia (Eyde, 1963). 


Family FAGACEAE Dum. 


The Fagaceae (oak family) are represented in the Nut 
Beds by at least two genera of fruits: Quercus, and 
Castanopsis. Two kinds of fagaceous foliage have been 
recovered from the Nut Beds (Manchester, 1981). In 
addition, two species of fagaceous wood have been 
recognized (Scott and Wheeler, 1992): Quercinium 
crystallifera, with anatomy corresponding both to ex- 
tant Lithocarpus and to evergreen species of Quercus, 
and Fagaceoxylon ostryopsoides. Fagaceoxylon Scott 
et Wheeler is an extinct genus with characters diag- 
nostic of fagaceous wood including aggregate rays, ор- 
posite to alternate intervascular pitting, mostly simple 
perforation plates, vasicentric tracheids and variable 
ray-vessel parenchyma pits. As noted by Scott and 
Wheeler (1982), however, the pore arrangement of Fa- 
gaceoxylon, i.e., diffuse porous with radial multiples 
and clusters arranged in “flame-like” tracts, is un- 
known in any living genus. 

The Fagaceae have an excellent fossil record in the 
Tertiary of the northern hemisphere (Daghlian and 
Crepet, 1983; Crepet and Nixon, 1989 a, b; Kvacek 
and Walther, 1989). The Nut Beds flora provides the 
oldest known fruit records of Quercus and Castanopsis. 


Genus CASTANOPSIS (D. Don) Spach. 


Castanopsis crepetii sp. nov. 
Plate 12, figures 6-11 


Etymology.— This species is named after William 
Crepet, recognizing his contributions to the North 
American fossil record of Fagaceae. 

Description.— Fruit a globose nut completely envel- 
oped by cupule, length (15)-39 mm, equatorial di- 
ameter (15)-36 mm; base and apex rounded, surface 
coarse; nutshell 1.8-2.0 mm thick, composed of iso- 
diametric angular sclereids 12-50 um in diameter ОГ 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 47 


ganized concentrically into islands of densely packed 
Cells among smaller cells, with occasional vascular 
bundles passing through the nutshell; cupule adnate to 
the nut, 1.8-2.0 mm thick, composed of similar, less 
densely packed cells. Locule cast rounded-conical, 
truncate basally, pointed apically, with meridional 
grooves defining broad-rounded ridges. 

Specimens.—Holotype: UF 9267, large perminer- 
alized fruit. Paratypes: UF 9268, USNM 354811, 
446079, fruits; UF 9597, 9598, locule casts. 

Discussion.—This species is represented by permi- 
eralized fruits and occasional locule casts. The ho- 
lotype isa large, fragmentary permineralized specimen 
showing the globose outer surface of the cupule (Pl. 
13, fig, 6), with portions of the cupule and nutshell 
removable to show the apically tapered, basally trun- 
Cate, locule cast with longitudinal grooves (Pl. 12, figs. 
7, 8). Thin sections of the wall clearly show the junction 
Of cupule with the nut (Pl. 12, figs. 9, 10). 

In the Fagaceae, some species of both Castanopsis 
and Lithocarpus have globose nuts with complete cu- 
Pular coverage and adnation between the nut and cu- 
Pule (Camus, 1952-54; Kaul, 1988, 1989). These gen- 
ста apparently overlap with one another, as summarized 
by Kaul (1989, р. 68): “Generic and infrageneric tax- 
onomy in the Lithocarpus—Castanopsis complex has a 
long history of instability (Forman, 1966). Some recent 
authors have accepted the two genera in their broad 
Senses, acknowledging some species with intermediate 
Character states." 

Castanopsis piriformis Hick. et A. Camus from An- 
Nam, southern Vietnam (Pl. 12, figs. 12-15) isan extant 
Species that resembles C. crepetii in size, globose shape 
Of the fruit, wall structure and locule morphology. 
Transverse sections of the endocarps of these two spe- 
Cles reveal remarkable similarity, both at the macro- 
Scopic and microscopic levels. In both the fossil and 
the modern species, adnation of cupule and nut is clear- 
ly evident (Pl. 12, figs. 9, 14) and the nutshell wall is 
Made up of more-or-less angular sclereids that are 
Srouped concentrically to form conspicuous “islands” 
85 viewed in transverse section (Pl. 12, figs. 11, 15). 

he anatomical correspondence confirms the identity 
i this fossil to the Fagaceae, and supports placement 
Within Castanopsis (providing that the generic assign- 
ment of C. piriformis is correct). Castanopsis also has 
a 800d fossil fruit record in Europe based upon species 
with small, angular nuts (Mai, 1989). 


Genus QUERCUS L. 


Quercus paleocarpa Manchester 
Plate 12, figures 16-18 


Quercus paleocarpa Manchester, 1976, p. 82, pl. 5, fig. 29. 


Description.— Acorn with prolate to ovate nut cov- 
ered А to У; from base to apex by a thick woody cupule; 
complete fruit length (nut with cupule) 22 mm, width 
15.5-21.8 mm; nut length (without cupule) 14.0-18.0, 
width 11.4-15.6 mm; nut base rounded, apex obtuse; 
relatively smooth, finely longitudinally striate; cupule 
bowl-shaped, height 9.0-10.4 mm, width 18-21 mm, 
with seven concentric exterior transverse ribs; the ribs 
decreasing in thickness from the base to the apical rim. 

Specimens :— Holotype: ОМІ Pb176, nut with 
abraded cupule. Hypotypes: USNM 414508, isolated 
cupule; USNM 312758 (Bones, 1979, pl. 6, fig. 9), 
complete, abraded nut. 

Discussion.—This species, representing the oldest 
known acorn, is represented only by a few specimens. 
The most complete specimen, with both the nut and 
cupule intact (Pl. 12, figs. 16, 17) was abraded prior to 
deposition, so that the arrangement of involucral scales 
on the outer surface of the cupule cannot be deter- 
mined. A well-preserved detached cupule (Pl. 12, fig. 
18) shows the outer surface in an unworn condition, 
revealing involucral scales arranged in concentric rings. 
I assume that each of these specimens represents the 
same species, although a larger population would be 
desirable to rule out the possibility that two different 
taxa are present. 

Although the cupules of extant species of Lithocar- 
pus and Quercus are often indistinguishable (Kaul, 
1988), the coarse, rather than papery, lamellae of the 
cupule and the limited extent to which the cupule cov- 
ers the nut suggest that the species belongs to Quercus 
rather than Lithocarpus. Cupules with the involucre 
arranged in concentric rings as in this fossil occur in 
the strictly Asian subgenus Cyclobalanopsis of Quer- 
сиз, as well as in Lithocarpus. 

Nixon (1989) noted that the cyclic arrangement of 
cupule scales/spines, which also occurs in Chrysolepis, 
Castanopsis, Castanea and Trigonobalanus, is appar- 
ently pleisiomorphic within the Fagaceae, with the im- 
bricate-spiral arrangement found in the derived forms 
of Quercus. If this interpretation is correct, then it is 
not surprising that Quercus paleocarpa, the oldest 
known Quercus fruit, should possess a cupule with cy- 
clically arranged scales. 


Family FLACOURTIACEAE DC. 


The Flacourtiaceae are a family of about 90 genera 
and about 875 species of shrubs, less often trees, that 
are chiefly tropical in distribution, but with a few that 
are temperate. The species described below, belonging 
to an extinct genus, is the only member of this family 
recognized from the Nut Beds. The extant genus On- 
coba, recognized in the London Clay flora (Reid and 


48 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Зтт 


Text-figure 11.—Saxifragispermum tetragonalis sp. nov., trans- 
verse section showing four-valved capsule with parietally attached 
seeds. 


Chandler, 1933) does not appear to be present in the 
Nut Beds collections. 


Genus SAXIFRAGISPERMUM Reid et Chandler 


Saxifragispermum tetragonalis sp. nov. 
Plate 13, figures 1-16; Text-figure 11 


Etymology.— Tetra (Gr = four) + gonia (Gr = angle, 
corner), referring to the four-angled shape of the fruit. 

Description.—Fruit an elongate-obovoid, tetracar- 
pellate, tetravalved loculicidal capsule, nearly circular 
to rounded-quadrangular in cross section; length 6.4— 
11.8, avg. 9.2 mm (SD=1.42, n=20), width 3.0-5.6, 
avg. 4.2 mm (SD=0.63 n=19); base more or less round- 
ed, sometimes with a small projection at the pedicel 
scar, apex pointed; placentation parietal, with four 
elongate placental ribs, one positioned medially on each 
of the four valves of the fruit and extending the full 
length of fruit; seeds numerous, six to eight radiating 
from any given level on the placenta, arranged verti- 
cally in rows about 12 high; interior of fruit solid with 
sinuously arranged fibers 5-18 um in diameter having 
a general longitudinal orientation, among which the 
seeds are embedded; fruit wall comprised of two layers: 
inner layer forming a thin lining (80-120 um thick) of 
longitudinally oriented fibers 12-30 um in diameter 
(this is the layer that forms the placental ribs), outer 
layer (150-200 um thick) formed of similar fibers ori- 
ented horizontally and periclinally; this outer layer 
showing lines of loculicidal dehiscence; pedicel scar 
with five apertures, one central, and four lateral, rep- 
resenting extensions of the placentae. Seeds ovoid, as- 
cending, rounded basally, tapered apically, 0.5-0.7 mm 
long, 0.35-0.45 mm wide; seed coat composed of an 
outer layer of mostly quadrangular cells about 15-20 
um wide producing small spine-like simple hairs 18- 
25 um long and an inner layer of transversely elongate 
hexagonal cells about 35-40 um wide and 12-14 um 
high. 

Specimens.—Holotype: UF 5236 (Bones, 1979, pl. 


4, fig. 4). Paratypes: UCMP 10614-10617, UF 6312, 
6471, 6473-6478, 6575, 6649-6653, 6972, 9897, 
USNM 353999, 354000 (21 specimens), 354001 (7 
specimens), 354002 (2 specimens), 354006-354008, 
354009 (7 specimens), 354010 (2 specimens), 354011, 
355469. 

Discussion. — This species is represented by numer- 
ous permineralized specimens and occasional casts. 
Part of the outer wall is often stripped away when the 
specimens are removed from the matrix, sometimes 
showing the rows of seeds (Pl. 13, figs. 3-5). Serial 
transverse sections reveal as many as eight seeds sur- 
rounding each placenta in a single slice (Pl. 13, figs. 7— 
9). The placentae are preserved in white silica and 
appear to have been hollow or parenchymatous (Pl. 
13, fig. 9). Scanning electron microscopy of fractured 
specimens illustrates details ofthe seeds and surround- 
ing fibers (Pl. 13, figs. 11-14). 

These valvate capsules resemble the fruits that Reid 
and Chandler (1933) named Saxifragispermum spi- 
nosissimum (emended diagnosis Chandler, 1961b) from 
the London Clay. Both have median longitudinal pa- 
rietal placentae with about a dozen rows of seeds on 
each placenta; in both taxa the entire cavity of the 
locule is virtually filled with fibers (Pl. 13, figs. 9, 11- 
13). Seeds of the London Clay fossil were originally 
described as spinose and small projections from the 
seed coat were figured by Reid and Chandler (1933, 
pl. 8, fig. 35). However, Chandler (1961b) redescribed 
the seeds more cautiously: “Testa apparently spines- 
cent (unless this is an effect of decay)”. Thin sections 
of the Nut beds specimens confirm the presence of 
small spines or spine-like trichomes on the seed coat 
(PI. 13, figs. 15, 16). 

Placentation and seed morphology indicate that 
Saxifragispermum is related to the Flacourtiaceae. Al- 
though the generic name suggest affinities to Saxifra- 
gaceae (Reid and Chandler, 1933), Chandler (1961a) 
reported that “Examination of more material and re- 
consideration of the whole evidence necessitates а cor- 
rection of the supposed relationship to Saxifragaceae. 
The single locule with its broad parietal placentae bear- 
ing several rows of seeds is unlike any known Saxifra- 
gaceae, a family which normally shows two or more 
locules. It is highly suggestive of Flacourtiaceae. Моге- 
over the succession of coats in the seed and the char“ 
acter of the chalaza are so closely comparable with the 
same features in Oncoba that the probability of rela- 
tionship with Flacourtiaceae is confirmed.” 

Although clearly congeneric with the London Clay 
fruits, the Nut Beds material can confidently be ге“ 
garded as a distinct species. The London Clay fruits 
are much wider in diameter, and have larger seeds. 5. 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 49 


Spinosissimum fruits range from 11.5 to 15.75 mm, 
whereas those of S. tetragonalis range only from 6.4— 
11.8 mm. The London Clay fossils range from four to 
five carpels, whereas the many Nut Beds specimens 
are all tetracarpellate. Carpolithus subfusiformis (Bow- 
erbank) Reid et Chandler (1933), which is known from 
tri- and tetracarpellate fruits, also shows similarity to 
and may, in fact, be conspecific with S. spinosissimum. 


Family HAMAMELIDACEAE R. Br. 


The Hamamelidaceae (witch hazel family), with 29 
extant genera and about 90 species, is divided into four 
Subfamilies: the Hamamelidoideae, Rhodoleioideae, 
Exbucklandioideae, and Liquidambaroideae (Endress, 
19893). The Liquidambaroideae are known to have 
been present in the Eocene based upon Liquidambar 
leaves in the Green River Formation, Utah (Mac- 
Ginitie, 1969), and fruits identified as Steinhauera Presl 
from the Geiseltal of Germany (Mai, 1976), but have 
Not been identified from the Clarno Formation. In- 
fructescences from the Clarno Nut Beds formerly iden- 
üfied as Altingia (Scott in Chandler, 1961a; Bones, 
1979) have been shown to represent Macginitiea (Pla- 
tanaceae; Manchester, 1986). The Hamamelidoideae 
are represented in the Nut Beds flora by the single fruit 
and seed species described below, and by inflorescences 
that may or may not represent the same taxon (Pl. 14, 
figs. 14—16). The oldest fossils with close affinities to 
the Hamamelidaceae are flowers with intact stamens 
ànd pollen from the Late Santonian or Early Campan- 
làn of Scania, Sweden (Endress and Friis, 1990). 


Genus FORTUNEARITES gen. nov. 


4 Etymology.— Fortunearites is named for its similar- 
ПУ to extant Fortunearia. 
Type species.— Fortunearites endressii sp. nov. 


Fortunearites endressii sp. nov. 
Plate 14, figures 1-13 


Etymology.— The species 15 named for Peter К. En- 
Tess, recognizing his important contributions in un- 

derstanding the systematics and phylogeny of Hama- 
Melidaceae. 

Description— Infructescence a spike, with a stout axis 
UP to 53 mm long, 2.4 mm in diameter, bearing nu- 
Merous closely packed sessile fruits; infructescence 
about 15 fruits in length with up to eight fruits en- 
countered at a given level in transverse section of the 
axis; fruit a bicarpellate, bilocular woody capsule, 
Quadrilaterally symmetrical, squarish in longitudinal 
View, elliptical in transverse section, with the long axis 
of ellipse parallel to the infructescence axis; capsule 5- 

mm long, 6.8-7.5 wide, 5.0-5.2 mm thick, fruit wall 


0.8-1.0 mm thick; locules more-or-less pyriform, flat- 
tened ventrally where they contact the septum, single- 
seeded; dehiscence of capsules apical by a split bisect- 
ing each of the locules through the major plane of 
symmetry. 

Internal molds of the seed coat obliquely ellipsoidal, 
asymmetrical, pointed apically, rounded basally, 5.8— 
6.2 mm long, 3.3-3.4 mm wide, 2.4-2.8 mm thick, 
length/width ratio 1.7-1.8; hilar scar not evident in 
these molds; surface of mold smooth except for a raised 
ribbon-like raphe band, 0.4-0.5 mm wide extending 
from the front to the back of the seed passing obliquely 
over the base in a u-shaped course with each limb 
extending У to 25 of the distance toward the seed apex. 

Specimens.— Holotype: OMSI Pb156 (and counter- 
part, USNM 354581), permineralized infructescence. 
Paratypes: UF 8741, permineralized infructescence; UF 
9258, 9259, locule casts; UF 6530 9330, USNM 
424632, 424651, internal molds of seed coat. 

Discussion.— This species is represented by two per- 
mineralized infructescences (e.g., Pl. 14, figs. 1, 2), a 
few locule casts (e.g., Pl. 14, figs. 6-9), and several 
endosperm casts (e.g., Pl. 14, figs. 10-13) from the Nut 
beds. Tangential sections of the infructescence inter- 
cept individual fruits transversely (Pl. 14, figs. 3, 4) 
showing the thick walls and bilocular construction of 
the capsules. Seeds are not preserved within the in- 
fructescences and it is likely that they had dispersed 
prior to deposition. Isolated seeds (actually internal 
molds of the seed coat, Pl. 14, figs. 10-13) correspond 
in shape and size to the locules of the infructescences. 
The capsule morphology matches that of extant genera 
in the subfamily Hamamelidoideae. Genera of this 
subfamily have one seed per carpel, which is ejected 
explosively at maturity. The large number of fruits on 
the axis, and seed characters discussed below, suggest 
affinity with extant Sinowilsonia and Fortunearia. 

Endress (1989b) indicated that seeds of extant Ha- 
mamelidoideae are structurally constrained by the ex- 
plosive dehiscence mechanism so that fruit and seed 
structure is extremely uniform throughout the subfam- 
ily, making it difficult to determine the affinities of 
fossil seeds down to the generic level., Subsequently, 
Grote (1989) found differences in hilar scar morphol- 
ogy among some of the extant genera that enabled 
recognition of Corylopsis seeds in the Eocene of Ten- 
nessee. In addition, I have observed differences in the 
course of the raphe that set Sinowilsonia and Fortu- 
nearia apart from other genera of the subfamily. In 
reviewing the seed anatomy of selected extant genera 
of Hamamelidaceae, Mohana Rao (1974) indicated that 
seeds of Hamamelis, Parrotia and Sinowilsonia have 
a raphe with vascular bundles running along one side 


50 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


5mm 


Text-figure 12.—Transverse sections of Hydrangea knowltonii sp. 
nov., showing deeply intruded parietal placentae and numerous seeds. 
A, Tricarpellate capsule. B, Bicarpellate capsule. 


of the seed ending at the chalaza. In dissections of 
modern seeds I found that the raphe bundles indeed 
run their course on only one side of the seed and ter- 
minate basally in Eustigma, Loropetalum, Matudaea, 
Parrotia, and Hamamelis. However, in Fortunearia 
sinensis Rehder et Wilson and Sinowilsonia henryi 
Hemsley these bundles curve over the base of the seed 
and extend apically on both sides of the seed. The 
strap-like ridge on the internal seed molds of Fortu- 
nearites (Pl. 14, figs. 10-13), representing the raphe, 
conforms to the morphology in Fortunearia and Si- 
nowilsonia. These two genera have been recognized as 
sister taxa on the basis of other characters and are 
placed with the more distantly related Eustigma in the 
subtribe Eustigmateae Harms (Endress, 1989a, c). 

Compact globose staminate inflorescences with ha- 
mamelidaceous pollen have also been recovered from 
the Nut Beds (Pl. 14, figs. 14-16, Pl. 15, figs. 1-5). 
These inflorescences are apparently unisexual with 
flowers having a pair of pistilodes, a whorl of ten sta- 
mens and staminodia, five narrow petals and five se- 
pals with a dense covering of stellate trichomes (Pl. 14, 
fig. 16). Only two specimens have been recovered (UF 
5567, 6109). Since they appear to belong to the Ha- 
mamelidoideae, it is possible that these inflorescences 
also belong to Fortunearites. The flowers share with 
Fortunearia and Sinowilsonia the presence of narrow 
petals, and conspicuously introrse anthers. On the oth- 
er hand, they differ from those of extant Fortunearia 
and Sinowilsonia in having ten stamens + staminodes 
per flower (Pl. 14, fig. 15) rather than five. Pollen from 
the stamens of the fossil (Pl. 15, figs. 1-5) is more 
similar to that of other genera such as Hamamelis and 
Corylopsis than to that of extant Fortunearia and Si- 
nowilsonia (Bogle and Philbrick, 1980). If these inflo- 
rescences indeed belong to the same species as the 
infructescence, then the case for treating Fortunearites 
as an extinct genus is strengthened. 


Family HYDRANGEACEAE Dumort. 


The Hydrangeaceae include 17 extant genera and 
about 170 species of trees, shrubs, lianas and herbs, 
predominately temperate and distributed mostly in the 
northern hemisphere. Morphological, chemical and 
chloroplast DNA (rbcL) characters indicate that Hy- 
drangeaceae are closely allied to the Cornaceae (Mor- 
gan and Soltis, 1993; Xiang et al., 1993). The family 
is represented in the Tertiary of Europe by fruits of 
Dichroa, Hydrangea, Schizophraqma and Itea (Mai, 
1985). The showy, usually tetrasepalous, sterile flowers 
of Hydrangea are easily identified (Pl. 15, figs. 6, 9) 
and have been recognized from many Tertiary local- 
ities in western North America. 


Genus HYDRANGEA L. 


Hydrangea knowltonii sp. nov. 
Plate 15, figures 7, 8; Plate 16, figures 1-15; 
Text-figure 12 


Etymology.—This species is named after the late 
Frank H. Knowlton, who was the first to recognize 
flowers of this genus in the Tertiary of North America. 

Description.—Infructescence a compound, trichot- 
omously branching dichasium at least 43 mm long; 
pedicels 1.5-2.7 mm long; fruits bicarpellate and tri- 
carpellate obovoid to subconical capsules. Fruit di- 
mensions from compression specimens: length 1.2-2.0 
mm, width 1.5-2.2 mm; base acute, apex truncate with 
an epigynous disk; exterior of fruit with about ten lon- 
gitudinal ribs; styles two to three, persistent, free, 0.6- 
0.8 mm long. 

Silicified fruits bicarpellate and tricarpellate multi- 
seeded thin-walled capsules, prolate, two- to three-lobed 
and correspondingly bi- and trisymmetrical; locule casts 
rounded apically and basally, height 2.4-4.0, avg. 32 
mm (SD=0.44, n=20), maximum width in bilobed 
specimens 3.2-4.2, avg. 3.8 mm (SD=0.34, n=13), 
maximum width in trilobed specimens 3.2-4.5, avg. 
4.1 mm (SD=0.53, n=5), width of individual lobes 
2.3-3.8, avg. 3.1 mm (SD=0.41, n=1 5); endocarp wall 
about 45 um thick, composed of transversely aligned 
fibers; apical end of the locule cast with a characteristic 
two- or three-angled wedge (silica-filled dehiscence 
opening) at the intersection of the two or three carpels; 
fruit unilocular, placentation deeply intruded parietal, 
almost axile, septa 90 um thick, bifurcating near the 
center of fruit and producing a placenta from each arm. 
Seeds numerous, tiny, winged, 625 um wide, 225-270 
um thick and 900 or more um long, with a peg-like 
hilum and a flattened, elliptical central body, encircled 
longitudinally by a wing; seed coat uniseriate, made 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 51 


up of rectangular cells 33-58 um in diameter, drawn 
Out laterally into a flange that forms the wing (typically 
180 um wide); embryo straight to sinuous. 

Specimens.—Holotype: UF 6445, locule cast. Para- 
types: OMSI Pb475, Pb697, infructescence compres- 
Sions; UCMP 10668, 10669, UF 6444, 6446, 8691- 
8699, 9384, 9385, 9423, 9883, USNM 312763 (Bones, 
1979, pl. 6, fig. 11), 312764 (Bones, 1979, pl. 6, fig. 
12), 355390, 355464 (3 specimens), 355651 (2 speci- 
mens) 355652 (13 specimens), 355658, locule casts. 

Discussion.— Hydrangea is represented in the Nut 

Beds by a few compression specimens from the leaf 
horizon at the base of Face 3 and by many isolated 
Chalcedony locule casts from the upper part of Face 3. 
The affinities of the compression fossils are readily 
apparent because the trichotomous mode of branching 
Of the infructescences (Pl. 15, fig. 7) and the morph- 
Ology of the fruits with their longitudinal ribbing, epig- 
ynous disk, and persistent styles (Pl. 15, fig. 8) precisely 
Match the extant genus (PI. 15, fig. 9). 
_ The isolated silicified fruits were more difficult to 
identify at first because surface details, including the 
disk and persistent styles, are not preserved. Instead, 
the specimens are preserved mainly as locule casts with 
only occasional details of the fruit wall remaining. The 
distinctive, transversely striate locule casts vary from 
two- to three-lobed, i.e., two to three-carpellate and 
havea conspicuous two- to three-armed apical “wedge” 
(PL 16, figs. 1, 3, 6) representing the dehiscence open- 
ing. Five tricarpellate (e.g., Pl. 16, figs. 1, 2) and 17 
bicarpellate specimens (e.g., Pl. 16, figs. 3, 4) were 
Observed. In addition, numerous specimens are rep- 
Tesented by detached unicarpellate lobes of the locule 
Cast, broken in the process of collecting. Although the 
fruit wall itself is usually not preserved, striations on 
the locule casts (Pl. 16, figs. 1-7, 9) indicate that the 
Surrounding endocarp was composed of transversely 
oriented fibers. The pericarp is partially preserved in 
Опе specimen (Pl. 16, fig. 9) showing that the thin 
endocarp is surrounded by a mesocarp composed 
Mostly of isodiametric cells but with widely spaced 
longitudinally oriented bands of fibers. The longitu- 
dinal fiber bands of the outer layer in the silicified 
Specimen correspond to those readily observed on the 
Impression specimens (Pl. 15, fig. 8). 

Due to the translucency of the chalcedony locule 
Casts, seeds can sometimes be observed within them 
45 tiny dark spots. Microscopic details of individual 
Seeds may be observed both in thin sections of the 
fruits (Pl. 16, figs. 12, 13) and in preparations made 
by dissolving the fruits in hydrofluoric acid (Pl. 16, 
figs, 14, 15). The seeds have a thick, peg-like hilum 
апа are winged. Serial transverse sections of the silic- 
¡fed Specimens show that the seeds were borne on 


deeply intruded placentae (Pl. 16, figs. 8, 10, 11), as 
illustrated in Text-fig. 12. 

In fruit shape and size, apical dehiscence, wall anat- 
omy, placentation, seed morphology, and in the vari- 
ability between two- and three-carpellate fruits, these 
specimens correspond precisely to extant Hydrangea. 
In her monograph of the genus, McClintock (1957) 
noted that variations in seed morphology (seeds cau- 
date or not, winged or not) are of systematic utility. 
Seeds with a circular wing surrounding the entire seed 
body are confined to Hydrangea, section Hydrangea, 
subsection Calyptranthe (Maxim.) McClintock, which 
has a single modern species, H. anamola D. Don, a 
widely distributed species extending from eastern 
Himalyaya of northern India to Korea and Japan (PI. 
16, fig. 16). 

Hydrangea is also known from other localities of the 
Clarno Formation (West Branch Creek, White Cliffs, 
Gonser Road) on the basis of conspicuous tetrasepal- 
ous Sterile flowers (Pl. 15, fig. 6) as well as by fruiting 
dichasia similar to that described above. Today, Hy- 
drangea includes about 23 modern species that are 
distributed from the Himalayas to Japan and Malesia, 
eastern North America, northern Central America and 
western South America. 


Family ICACINACEAE Miers 


The Icacinaceae are a family of trees, shrubs and 
lianas, mostly of tropical distribution, with about 60 
extant genera and 320 species. Fruits typically are uni- 
locular, single-seeded drupes. The family is tradition- 
ally divided into four tribes (Engler, 1897, 1964): the 
Icacineae (mostly trees, shrubs, seldom climbers, ves- 
sel perforations typically scalariform), the Iodeae (li- 
anas with dioeceous flowers and vessels with simple 
perforations), the Sarcostigmateae (lianas, fruit locule 
weakly wrinkled) and the Phytocreneae (lianas, fruit 
locule warty or spiny). The general validity of these 
tribes was supported by studies of vegetative anatomy 
(Bailey and Howard, 1941; Howard, 1942), although 
there is need for refinement. The family is common in 
the early Tertiary of western North America, Europe 
and Asia, as reviewed by Tanai (1990): The family is 
represented in the Nut Beds by five genera and eight 
species: Comicilabium, Iodes (two spp.), Iodicarpa (two 
spp., Palaeophytocrene (two spp.) Pyrenacantha. 
These genera belong to the tribes Iodeae and Phyto- 
creneae, and thus probably represent lianas. 


Tribe IODEAE Engler 
Genus IODES Blume 


Iodes multireticulata Reid et Chandler 
Plate 17, figures 1-8 


52 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Description.—Endocarp bilaterally symmetrical, 
unilocular, single-seeded, broadly oval or ovate, lat- 
erally compressed in the plane of symmetry, lenticular 
in transverse section, pointed apically, rounded ba- 
sally; length 3.5-4.7, avg. 4.0 mm (SD = 0.51, n = 4), 
width 6.4-8.0, avg. 7.1 mm (SD = 0.66, n = 4), thick- 
ness 8.0—10.0, avg. 9.3 mm (SD = 0.96, п = 4); exterior 
of endocarp with reticulate pattern of sharp ridges that 
delimit numerous shallow polygonal depressions, giv- 
ing the appearance of areolate venation with occasional 
simple freely ending veinlets; with a keel surrounding 
the fruit in the plane of symmetry, this keel thicker on 
one of the lateral margins than on the other, containing 
the funicular canal which runs from the base to the 
apex; endocarp wall 450 um thick, formed of dense 
sclerenchyma (details of cell shape not known). Locule 
cast similar in shape to the endocarp, but without sharp 
ridges, having 40—50 shallow depressions on each side, 
corresponding to the reticulations of the endocarp; 
length 6.0-11.7, avg. 7.6 mm (SD=1.7, n=11), width 
4.5-9.7, avg. 6.1 mm (SD=1.42, n=11), thickness 1.2— 
4.4, avg. 2.8 mm (SD=0.88, n=10); locule cast surface 
dotted with closely and regularly spaced fine punctae 
(15-21 um in diameter), about 12-16 per square mm; 
seed anatropous, pendulous, with apical hilum, basal 
circular chalaza (0.8-1.0 mm in diameter) and the ra- 
phe running longitudinally across one lateral face; sur- 
face of seed coat composed of rows of square to rect- 
angular cells, 20-30 um in diameter. 

Specimens.—UCMP 10702, endocarp cast; UF 6458, 
USNM 446053, endocarp mold and locule cast; OMSI 
Pb180, UCMP 10714, UF 8592-8595, 9805, USNM 
354779, 354781, 355034, 355526, 422394, 424647, 
locule casts; HU 59974, one endocarp valve. Holotype: 
BM у22589, locule cast from the London Clay. 

Discussion. — This species is represented in the Nut 
Beds collections primarily by chalcedony locule casts, 
which are readily recognized by their faceted surfaces 
(Pl. 17, fig. 5). Close examination reveals regularly 
spaced fine punctae on the surface (Pl. 17, figs. 7, 8), 
indicating that the inner lining of the endocarp was 
papillate, as in extant Todes (e.g., I. cirrhosa, Pl. 19, 
fig. 18). The endocarp itself is usually not preserved, 
but is represented by a space in the rock between the 
locule cast and the sedimentary mold of the endocarp 
exterior. The conspicuous reticulate sculpture of the 
endocarp surface may be seen on the surface of rare 
endocarp casts (Pl. 17, figs. 1, 2) and as an impression 
in the sedimentary molds of the endocarp (Pl. 17, figs. 
3, 4). In some of the locule casts, a thin layer of chal- 
cedony has fractured away to expose cells of the seed 
coat, and in some cases morphological features of the 
seed, including the hilum, raphe and chalaza, are readi- 
ly apparent on the locule cast surface (Pl. 17, fig. 5). 

The Nut Beds specimens conform closely in all de- 


tails of their preservation to the fossils designated odes 
multireticulata Reid et Chandler (1933) from the Lon- 
don Clay flora (Reid and Chandler 1933), and I have 
found no morphological or anatomical features to dis- 
tinguish them as a separate species. Papillae lining the 
locule are reported by Reid and Chandler to be 0.05 
mm in diameter, whereas those on the Clarno speci- 
mens measure 0.16 to 0.21 mm. This difference may 
be attributed to measurements being taken from near 
the apex, rather than at the basal neck of the balloon- 
like papillae. 

Placement of this species in the Icacinaceae is jus- 
tified by many features including the bilaterally sym- 
metrical, unilocular fruits, marginal funicular canal, 
and the raphe running along one of the flattened faces 
(Reid and Chandler, 1933). Assignment to the tribe 
Iodeae is supported by the combination of reticulate 
endocarp sculpture and papillate locule lining. The fos- 
sil endocarps are somewhat smaller than endocarps of 
the living species of Jodes that I have examined. Other 
genera with similar endocarps that are faceted and have 
papillate locules include Hosiea Hemsley and E. Wil- 
son, with two species in China and Japan (see Mai and 
Walther, 1978; Mai, 1987 for fossil reports), and Nat- 
siatum Buch.-Ham. ex Arn. with one species from east- 
ern Himalaya to southeast Asia. Chandler (1962) at- 
tributed an Eocene British species to Natsiatum on the 
basis of comparison with N. sinense Oliv.; however, 
this extant species is now treated as Hosiea sinensis 
(Oliv. Hemsl., and Chandler’s species was subse- 
quently placed in Hosiea (Takhtajan, 1966; Mai and 
Walther, 1978). 

Seeking characters by which to distinguish the en- 
docarps of Hosiea and Natsiatum from those of Todes, 
I examined specimens of Hosiea japonica, H. sinensis 
and Natsiatum herpeticum Ham. at the US National 
and Harvard herbaria. Transverse fractures revealed 
that endocarps of each species have a papillate locule 
lining like that of /odes, but lack a funicular canal 
within the endocarp (presumably this runs along the 
outside of the endocarp). By contrast, transverse sec- 
tions of Jodes endocarps clearly showed the funicular 
canal within the endocarp along one of the lateral mar- 
gins in the plane of symmetry. Although the ргезег- 
vation ofthe fossil does not show the funicular position 
unequivocally, it is interpreted to have been within the 
endocarp, supporting the placement in extant 10465. 
The position ofthe funicle in extant Natsiatum remains 
to be determined. Jodes, which ranges today from In- 
domalaya to Africa and Madagascar, is distinguished 
from other similar genera of the family by its opposite 
leaves, a condition that cannot be assessed from the 
isolated fossil fruits. 

Iodes, and/or closely related genera, are common in 
the Eocene of both Europe and North America. In 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 53 


Summarizing records from the Eocene of England, 
Chandler (1962) recognized about five species of odes 
and one species assigned to Natsiatum, with species 
distinguished on the basis of differences in overall shape, 
number and prominence of depressions on Ше locule 
Cast surface, presence/absence of stylar horns, and the 
Spacing and size of papillae lining the locule. Using 
Similar criteria, Knobloch and Mai (1986) provided a 
Morphological key to six European Tertiary fruit spe- 
Ges assigned to Годез. Subsequently, Mai (1987) rec- 
ognized two species of Hosiea and one of Iodes from 
the Paleocene of Germany. Collinson (1988) lists Nat- 
Siatum among fruits from the Middle Eocene Messel 
flora of Germany, an example of which is figured by 
Schaarschmidt (1988, p. 42). Similar endocarps are 
Preserved as impressions at other localities of the Clar- 
по Formation and in the middle Eocene Bridger For- 
mation of southwestern Wyoming (Blue Rim site, UF 
loc. 15761; S. R. Manchester, pers. obs.). 


Iodes chandlerae sp. nov. 
Plate 17, figures 9-16 


Etymology.— This species is named in memory of 

arjorie E.J. Chandler, recognizing the importance of 
her contributions to the study of fossil Icacinaceae and 
Other plant families. 

Description.— Locule cast more or less ovoid, bilat- 
егаПу symmetrical, somewhat compressed in the plane 
of Symmetry, elliptical in cross section; length 7.2-9.5, 
аур. 8.4 mm (SD=0.99, n=8), width 6.2-7.2, avg. 6.9 
mm (SD=0.37, n=8), thickness 4.3-5.6, avg. 5.2 mm 
(SD=0. 59, n=8); base truncate, apex asymmetric, 
Pointed; a keel or prominent ridge follows the plane 
of Symmetry from base to apex; in addition a promi- 
Dent ridge runs medially across the lower half of both 
longitudinal faces; the ridges intersect at right angles 
Nor the basal truncation; surface on some specimens 
faintly faceted and dotted with fine punctae. Seed one 
Per locule, surface of seed coat composed of rows of 
Square to rectangular cells 20-30 um in diameter ar- 
Tanged in rows. 

Specimens.—Holotype: USNM 424743. Paratypes: 
UCMP 10650, 10651, UF 9602, USNM 312760 

Bones, 1979, pl. 4, fig. 8), 355408 (3 specimens), 
355486 (2 specimens), 355402, 422383, 424740- 
424743, 

Discussion.— Locule casts of I. chandlerae differ from 
those of I. multireticulata by their more inflated, thick- 
°т appearance, less obvious surface depressions and 
truncate base. The affinities were unclear until a few 
Specimens showing the characteristic punctae were 
found, indicating a fruit with a papillate locule. In one 
9f the locule casts a layer was broken away, revealing 
а seed cast inside with details of the raphe and chalaza, 
and features of the seed coat surface (Pl. 17, figs. 9, 


10). The truncate base with a cross-like pattern of four 
intersecting meridional ribs (Pl. 17, figs. 14—16) some- 
times occurs in extant Jodes and is also seen in the 
icacinaceous fossil, Palaeophytocrene pseudopersica (Pl. 
20, fig. 10). The distribution and size of punctae (Pl. 
17, figs. 11, 12) are similar to that of Jodes multireti- 
culata. Because the species is known only from chal- 
cedony locule casts, external features of the endocarp 
remain unknown. 


Genus IODICARPA gen. nov. 


Etymology. — Iodeae (an extant tribe of Icacinaceae) 
+ karpos (Gr = fruit). 

Generic Diagnosis.—Endocarp ellipsoidal, bilater- 
ally symmetrical, compressed in the plane of sym- 
metry, length 26—56 mm, width 20—35 mm, thickness 
9-2] mm, unilocular, single-seeded, endocarp two- 
valved with dehiscence along the plane of symmetry; 
funicular canal passing from base to apex through the 
endocarp wall on one side in the plane of symmetry; 
endocarp wall thick (2-4 mm), composed of interlock- 
ing digitate cells with numerous intracellular spaces; 
locule smooth, lined by rounded balloon-shaped pa- 
pillae. 

Type species.— Iodicarpa ampla sp. nov. 

Discussion.— Iodicarpa is represented by two species 
in the Nut Beds assemblage. The fruits are easily dis- 
tinguished from those of odes by their large size and 
thick endocarp wall, but resemble Jodes in general form, 
endocarp wall composed of digitate cells and papillate 
locule lining (Pl. 19, fig. 18). 

It was considered that these fossils might be placed 
in the genus Icacinicarya Reid et Chandler. Icacini- 
carya was established to accommodate fossil endo- 
carps and seeds with characters diagnostic ofthe family 
Icacinaceae, but not assignable to a particular modern 
genus or section (tribe) within the family (Reid and 
Chandler, 1933). However, the Nut Beds fossils clearly 
conform to the tribe Iodeae, and differ from the Lon- 
don Clay species of Icacinicarya. Of the 12 species 
attributed to Icacinicarya by Reid and Chandler (1933), 
I. platycarpa (the first species that they described) is 
hereby designated the type species. The locule of 7. 
platycarpa is faceted, the locule lining is not papillate 
and the endocarp is formed of small cells **with no 
appearance of digitation, although some of the cells 
show sinuous outlines" (Reid and Chandler, 1933, p. 
345). Iodicarpa does not appear be represented among 
the London Clay fossils. 


Iodicarpa ampla sp. nov. 
Plate 18, figures 1-6, 11 


Etymology.— Ampla (L — large) referring to the large 
size of these fruits relative to those of Iodes. 


54 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Description.—Endocarp ellipsoidal, bilaterally sym- 
metrical, compressed in the plane of symmetry, with 
a width/thickness ratio of about 1.6, unilocular, single- 
seeded, endocarp length 26-35, avg. 31 mm (SD=2.66, 
n=12), width 21.0-30.4, аур. 26.0 mm, (SD=3.27, 
n=12), thickness 10.0-17.0, avg. 13.6 mm (SD=1.98, 
n=12), outer surface (rarely preserved) with a network 
of sharp ridges defining numerous polygonal areas in 
a pattern resembling areolate venation; worn endocarp 
surface rugulate, with rugulae mostly oriented longi- 
tudinally; endocarp two-valved with plane of splitting 
along the plane of symmetry; funicular canal passing 
from base to apex through the endocarp wall on one 
side in the plane of symmetry; endocarp wall 4.2 mm 
thick, composed of three anatomical layers: the outer 
Y made up of interlocking thick-walled anticlinally 
elongate cells with sinuous margins, 70-150 um long 
and 50-63 um wide; middle Y composed of similar 
elongate sinuous-walled cells mostly oriented longi- 
tudinally; inner У composed of similar but smaller 
interlocking cells; locule lined by rounded balloon- 
shaped papillae, 20-33 um wide, 27-33 um long. 

Specimens.—Holotype: USNM 354788, perminer- 
alized endocarp. Paratypes: OMSI Pb245, UF 8596, 
8618, UM 29940, USNM 354782, 355269, 446055, 
446085, endocarps; USNM 354159, 354198, 354241, 
354787, 354789, 354790, 354794, 354797, 354801, 
354802, 354804, 354806, 354808, 354813, 354814, 
354818, 354820, 354821, locule casts. 

Discussion.—This species is represented by permi- 
neralized endocarps and chalcedony locule casts. Be- 
cause of the way in which the carbonized or incom- 
pletely permineralized endocarps break out ofthe hard 
matrix with part ofthe wall remaining attached to the 
mold, the outermost surface of the endocarp is only 
rarely visible. The reticulate ridge pattern of the true 
endocarp surface is best seen as an impression surface 
on the sedimentary mold from which a carbonized 
endocarp has been removed (Р1. 18, fig. 3). Transverse 
sections of permineralized endocarps reveal the thick- 
ness and bivalved construction of the endocarp wall 
and the position of the funicular canal (Pl. 18, fig. 6). 
Thin sections show sinuous-walled cells making up the 
endocarp wall and papillae lining the locule (Pl. 18, 
fig. 11). 

This species corresponds to the Icacinaceae in the 
presence of two endocarp valves, one locule, one seed 
and with the funicle running through the thickness of 
the endocarp wall on one of the lateral margins, in the 
plane of bisymmetry (РІ. 18, fig. 6). Iodicarpa ampla 
resembles the extant genus /odes in the course of the 
funicle, the papillate locule lining and the reticulate 
ridge pattern of the endocarp surface. However, the 
fossil fruits are much larger and have thicker walls than 


those of any modern species of the /odes, and have a 
smooth, rather than faceted, locule. 


Iodicarpa lenticularis sp. nov. 
Plate 18, figures 7-10, 12-14 


Etymology.— Lentis (L = lens-shaped), referring to 
the endocarp and locule cast morphology. 

Description.— Endocarp ellipsoidal, bilaterally sym- 
metrical, compressed in the plane of symmetry, len- 
ticular in cross section, unilocular, single-seeded, width/ 
thickness ratio 1.9—2.2; endocarp length 31.6-56.1, avg. 
44.0 mm (SD=9.2, п=7), width 20.0-35.0, avg. 30.1 
mm (SD-5.2, п=8), thickness 9.4-20.5, avg. 4.9 mm 
(SD=33, n=8) (smaller one, UF 9745, 24 mm long, 
21.4 mm wide, 12.5 mm thick), exterior of endocarp 
relatively smooth; endocarp two-valved with dehis- 
cence along the plane of symmetry; funicular strand 
running in a conspicuous canal along a lateral margin 
of the endocarp within the wall; endocarp wall 2.0-2.5 
mm thick, composed of three anatomical layers: the 
outer % composed of interlocking thick walled anti- 
clinally elongate cells with sinuous margins, 60-220, 
avg. 130 um long and 30-65, avg. 45 um wide; middle 
А composed of similar elongate sinuous cells mostly 
oriented longitudinally; inner У composed of smaller 
isodiametric cells, 10-15, avg. 13 um in diameter; loc- 
ule lined by rounded balloon-shaped papillae, 18—25, 
avg. 21 um wide, 18—30, avg. 20 um long; seed coat 
uniseriate, composed of thin walled polygonal cells, 
15—25, avg. 20 um in diameter. 

Specimens.— Holotype: USNM 354816, perminer- 
alized endocarp. Paratypes: UF 8585-8591, 8614, 
8615, 8616, 9745, UM 29943, USNM 354816, 354796, 
354833, 354834, 354842, 354850, 354979, 354985, 
endocarps; USNM 354793, 354795, 354852, locule 
casts. 

Discussion.—A locule cast of this species was illus- 
trated by Scott (1954, pl. 16, fig. 31), but not named; 
only one specimen was known at that time. Specimens 
are preserved variously as isolated chalcedony locule 
casts, individual endocarp valve casts, and as complete 
carbonized or permineralized endocarps. Sections of 
permineralized endocarps reveal that the wall is com- 
posed of elongate cells with sinuous walls and show 
the conspicuously papillate locule lining (Pl. 18, figs- 
12-14). These anatomical features, along with the two- 
valved, unilocular, single-seeded endocarps, confirm 
the position of this species as a member of the Icaci- 


naceae. 

Anatomically, Ше endocarp of Jodicarpa lenticularis 
is virtually identical to that of 1. amplum. However 
I. lenticularis is readily distinguished by its greater 
length and commonly more compressed form, givin£ 
a greater width to thickness ratio, and by its smooth, 
rather than sculptured, endocarp surface. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 55 


Genus Comicilabium gen. nov. 


Etymology.— Comicus (L = funny) + labium (L = 
lip), referring to the shape of the funicular bulge. 
Type species.—Comicilabium atkinsii sp. nov. 


Comicilabium atkinsii sp. nov. 
Plate 19, figures 1-9 


Etymology.—This species is named for Alexander 
Atkins, in recognition of his invaluable assistance with 
field work at the Nut beds. 

Description.—Endocarp subglobose, bilaterally sym- 
metrical, unilocular, single-seeded, somewhat com- 
Pressed in the plane of symmetry; length 18.2-30.2, 
avg. 29.0 mm (SD=8.49, n=7); width 24.0-42.4, avg. 
30.9 mm (SD=7.02, n=7); thickness 21.0-30.2, avg. 
25.0 mm (SD=3.35, n=6); with about five faint lon- 
gitudinal grooves on both faces, radiating from a point 
at the base and extending toward the apical bulge; a 
fine groove running along the surface in the plane of 
bisymmetry disappearing over the apical bulge; apical 
bulge forming a conspicuous lip with a transverse slit 
8-9 mm wide perpendicular to the plane of bisym- 
Metry; funicular canal running longitudinally within 
the endocarp wall in the plane of symmetry on one 
Side, turning inward at the apical bulge; endocarp wall 
2-4 mm thick with a compact structure made up in- 
terlocking (jig-saw puzzle-like) digitate cells; cells of 
the outer part elongated with long axes anticlinal, 25— 
100 um in diameter; those of the inner part of the 
endocarp elongate with long axes horizontal/periclinal, 
10-12 um in diameter, typically 75 um long; locule 
lined by short bubble-like papillae 12-15 um in di- 
ameter; locule cast smooth, rounded basally, tapering 
to a point apically, with a keel along one side in the 
Plane of symmetry. 

Specimens. —Holotype: USNM 354783. Paratypes: 
UF 8596, OMSI Pb245, UM 29941, 66136, USNM 
354782, 355269, 446055, 446084, endocarps; USNM 
354159, 354198, 354241, 354787, 354790, 354794, 
354797. 354801, 354802, 354804-354806, 354808, 
35481 3, 354814, 354818, 354820, 354821, locule casts. 

Discussion.— This species is represented by endo- 
Carp casts, permineralized endocarps, and locule casts. 
It is conspicuous by the prominent apical placental 
bulge Where the funicle curves into the locule (Pl. 19, 
figs. 1,2, 5), and by the longitudinally oriented grooves 
9n the endocarp surface. A surface groove in the plane 
Of bisymmetry (Pl. 19, figs. 2-4) indicates that the fruits 
Were two-valved. Sectioned permineralized endocarps 
reveal a thick endocarp wall (Pl. 19, figs. 7, 8), a single 
Seed with paired subplanar cotyledons (Pl. 19, fig. 7), 
апа papillae lining the locule (Pl. 19, fig. 9). 

The combination of a unilocular endocarp, having 


the funicle on one side in the plane of symmetry, in- 
terlocking digitate cells and papillate locule lining con- 
firms affinities with the Icacinaceae. The papillate loc- 
ule lining is a condition found sporadically in the tribes 
Iodeae and Phytocrenae, and is seen in other icacin- 
aceous genera from the Nut Beds including Jodes (Pl. 
17, fig. 8), Iodicarpa (Pl. 18, figs. 12, 14), and Palaeo- 
phytocrene (Pl. 20, fig. 5). Because I have not found a 
modern genus with precisely corresponding morphol- 
ogy, I conclude that this species represents and extinct 
genus, for which the name Comicilabium is estab- 
lished. Comicilabium differs from Iodicarpa by the large 
funicular bulge, the straight meridional furrows, and 
its greater thickness measured across the plane of sym- 
metry. In addition, Comicilabium atkinsii differs from 
the Clarno species of /odicarpa in having larger, shorter 
papillae. 


Tribe PHYTOCRENEAE Engler 
Genus PALAEOPHYTOCRENE Reid et Chandler 


The genus Palaeophytocrene was established by Reid 
and Chandler (1933) to accommodate fossil fruits with 
tuberculate endocarps similar to those occurring in the 
extant tribe Phytocreneae of the Icacinaceae. These 
fossils are easily recognized by the prominent pits (tu- 
bercles) on the surface of endocarps and locule casts, 
and are relatively common in the Nut Beds (Scott, 
1954). In the distribution and nature of tubercles on 
the surface of the endocarp, the fossil genus resembles 
modern Phytocrene Wall (Pl. 19, fig. 16). As Reid and 
Chandler (1933) observed, however, the tubercles of 
Palaeophytocrene extend well into the locule whereas 
those of Phytocrene protrude only slighty, 1f at all, into 
the locule (Pl. 19, fig. 17). Other extant genera with 
tuberculate endocarps include Miquelia Meissn., Chla- 
mydocarya Baill., and Pyrenacantha Hook. These gen- 
era occur today in the Old World tropics. Detailed 
anatomical studies of extant fruits of the Phytocreneae 
have not been published, aside from the observations 
of Reid and Chandler (1933), and the functional sig- 
nificance of the tubercles (ог “сапа!5”, Scott, 1954) 
remains unknown. In contrast to Jodes, Comicilabium, 
and the /odicarpa species treated above, Palaeophy- 
tocrene, Phytocrene and Pyrenacantha do not show a 
funicle in cross sections of the endocarp. 

Palaeophytocrene is now known from many Paleo- 
gene localities in the northern hemisphere, including 
the Paleocene of North Dakota and Wyoming (Crane 
et al., 1990), the early Eocene Chalk Bluffs flora of 
California (MacGinitie, 1969, p. 132), the middle Eo- 
cene of Messel, Germany (Collinson, 1988), Republic 
Washington (W. Wehr, pers. comm.), and Bonanza, 
Utah (S. R. Manchester, pers. obs.), the Oligocene Gray 


56 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Butte, (М. A. Ashwill, pers. comm.), Dugout Gulch 
and Willamette (S. R. Manchester, pers. obs.) floras of 
Oregon, as well as from the London Clay flora (Reid 
and Chandler, 1933; Collinson, 1983). 

Scott (1954) recognized three species of Palaeophy- 
tocrene from the Nut Beds: Р. foveolata Reid et Chan- 
Чет, P. pseudopersica Scott and P. hancockii Scott, 
based upon six, three and one specimens, respectively. 
A much larger collection of Palaeophytocrene endo- 
carps and locule casts is now available, leading to a 
revised interpretation of this complex. I adopt only 
two of Scott’s Palaeophytocrene species for the Nut 
Beds material: P. hancockii and P. pseudopersica. Of 
the three specimens attributed by Scott (1954) to the 
London Clay species P. foveolata Reid et Chandler, 
one (his pl. 16, fig. 3) is treated here as Palaeophyto- 
crene pseudopersica, and two (pl. 16, figs. 1, 2) belong 
to Pyrenacantha occidentalis sp. nov. 


Palaeophytocrene hancockii Scott emend. 
Plate 20, figures 1-5 


Palaeophytocrene hancockii Scott 1954, p. 81, pl. 16, figs. 6, 7. 


Description.—Endocarp large, ovoid, bilaterally 
symmetrical, unilocular, single-seeded, somewhat 
compressed in the plane of symmetry, length 38.5- 
85.0, avg. 64.6 mm (SD=12.64, n=12), width 26.9- 
50.0, аур. 37.2 mm (SD=6.53, n=11), thickness 20.0- 
35.0, аур. 28.4 mm (50-4.56, п-9); outer surface 
covered by numerous pits, about 9 rows wide and 13- 
14 high, adjacent pits sometimes coalesced; endocarp 
wall 2.0-2.3 mm thick, composed of densely packed 
interlocking digitate cells, up to 200 um long and 30- 
80 um wide, arranged in layers parallel to the locule 
surface; tubercles (corresponding to the surface pits) 
about 2.2 mm in diameter, protruding 2.0-3.5 mm 
into the locule, straight-walled (cylindrical, sometimes 
bulbous-ended); locule finely papillate, papillae 10-15 
um high and wide; funicle not present in the endocarp. 

Specimens.—Holotype: UM 29933 (Scott, 1954, pl. 
16, figs. 6, 7). Hypotypes: USNM 312751 (Bones, 1979, 
pl. 4, fig. 6), UCMP 10715, 10716, UF 8603, 8604, 
8612, 9567, USNM 354420, 354155. 

Discussion.— Palaeophytocrene hancockii includes 
the largest of all fruits known from the Nut Beds. Scott 
(1954) distinguished this species from other species of 
Palaeophytocrene on the basis of its large size, and 
because, in proportion to its size, the tubercles protrude 
only a short distance into the locule by comparison 
with the other species of this genus. His description 
was based on a single specimen that had been sectioned 
transversely to show the endocarp wall, but in which 
the number and arrangement of pits are obscured. There 
are about 15 specimens now, including specimens that 


clearly show the number and arrangement of pits (PI. 
20, figs. 1, 2). Thin sections show the structure of the 
tubercles projecting into the locule (Pl. 20, figs. 3, 4), 
and the anatomy of the endocarp wall, including pa- 
pillae lining the locule (Pl. 20, fig. 5). 


Palaeophytocrene pseudopersica Scott emend. 
Plate 20, figures 6-13 
Palaeophytocrene pseudopersica Scott 1954, p. 80, pl. 16, figs. 4, 5. 


Palaeophytocrene foveolata Reid and Chandler. Scott 1954, pl. 16, 
fig. 3 only. 


Description.—Endocarp unilocular, single-seeded, 
oval to ovate, flattened or slightly inflated in the plane 
of bisymmetry, length 14.3-39.0, avg. 27.4 mm 
(SD=7.8, n=8), width 13.0-28.5, avg. 21.4 mm 
(SD=5.3, n=8), thickness 8.8-16.5, avg. 10.9 mm 
(SD=2.6, n=8); outer surface covered by numerous 
pits, eight to 14 pits wide, ca. 19 pits high (ca. 40—80 
pits on each half of the endocarp), adjacent pits some- 
times coalesced; endocarp wall 1.5-2.3 mm thick, 
composed of densely packed interlocking digitate cells; 
these cells up to 300 wm long and 20—50 um wide, 
oriented in layers parallel to the locule surface; hollow 
tubercles (corresponding to the surface pits) 1.2-2.4 
mm in diameter, protruding 1.7-3.1, avg. 2.2 mm into 
the locule, straight-walled (cylindrical), sometimes 
conical; locule lining, including that over the tubercles, 
finely papillate; papillae short, rounded 10-15 um high 
and wide, closely spaced. Seed not seen, but possible 
fragments of very thin seed coat observed in sectioned 
specimens. 

Specimens.—Holotype: UM 29931. Hypotypes: 
OMSI Pb178, Pb179, Pb181, Pb182, Pb247, UF 8597, 
8599, 8602, 8605-8610, 8613, 9568, 9569, UM 29928, 
29929, 29931, 29932. 

Discussion.—Scott (1954) described this species 10 
accommodate Clarno fossils that he noted were similar 
to the London Clay species, P. ambigua Reid and 
Chandler. The species is represented by abundant loc- 
ule casts (Pl. 20, figs. 12, 13) and permineralized en- 
docarps (Pl. 20, figs. 6-10) that range widely in size. 
The above description incorporates new anatomical 
data observed in thin sections (e.g., Pl. 20, fig. 11). 

As far as I can determine, there is not a very clear 
distinction between P. pseudopersica and P. hancockii, 
and the practice has simply been to place large speci- 
mens (38-85 mm long) in P. hancockii and smaller 
specimens, usually less than 40 mm long, in P. pseu- 
dopersica. The variation in size among specimens O 
Palaeophytocrene from the Nut beds is great. I ex- 
cluded two of the smallest specimens from the above 
description (UF 8609, 8610, which are 7.8, 11.8 mm 
long, 7.0, 9.0 mm wide, and 5.0, 6.0 mm thick), be- 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 57 


Cause they might be immature, or could represent an- 
Other species. The variation is more or less continuous, 
however, and, if more than one species is represented, 
It is difficult to draw distinct boundaries between them. 
Scott noted that in proportion to its size, the tubercles 
of P. hancockii protrude only a short distance into the 
locule in comparison with P. pseudopersica. However, 
the same phenomenon is noted in comparing succes- 
sively larger specimens of P. pseudopersica. Evidently, 
the projections do not increase in thickness in direct 
Proportion with increased fruit size in this genus. I 
have chosen to maintain two of the species established 
by Scott (1954), while recognizing that the large col- 
lections of this genus require more thorough investi- 
gation to determine the number of species represented. 
In order to understand better the significance of the 
Size variability among Palaeophytocrene specimens 
from the Nut Beds, it would be desirable to obtain 
Information on the intraspecific and ontogenetic size 
Variability among related extant genera such as Mi- 
quelia, Chlamydocarya and Pyrenacantha. 


Genus PYRENACANTHA Hooker 


Pyrenacantha occidentalis sp. nov. 
Plate 19, figures 10-14 


Р alaeophytocrene foveolata Reid et Chandler. Scott 1954, pl. 16, fig. 
1, only. 


Etymology.— Occidens (L = of the west) referring to 
Ше western occurrence of this species, which contrasts 
with the distribution of extant species in the eastern 
hemisphere. 

i Description. — Епдосагр ovoid, bilaterally symmet- 
пса], slightly compressed in the plane of symmetry, 
unilocular, single-seeded; length 25.0-33.0, avg. 29.9 
mm (80-2.53, n=8), width 15.7-30.0, avg. 22.7 mm 
(SD=4.18, п=7), thickness 11.0-25.3, аур. 18.3 mm 
(SD=4.32, n=7), surface dotted with numerous small 
Pits about 1 mm in diameter, spaced 1-2 mm apart, 
12-14 rows of pits wide, 16-19 pits high; pits corre- 
SPonding to elongate vertical-sided, cylindrical, tuber- 
cles which project 4.5-5.2 mm into the locule (pro- 
truding more than У of the distance across the locule), 
endocarp wall 1.0 mm thick, uniform, without funic- 
Шаг canal; cells of the endocarp elongate, sinuous. Loc- 
ule lining not papillate, seed conforming to the locule 
Projections. 

Specimens.—Holotype: UCMP 10708. Paratypes: 
ОМ] Pb170, Pb202, UCMP 10706, UF 8617, UM 
29927 (Scott, 1954, pl. 16, fig. 1), USNM 354767, 
354987 (and another portion of the same specimen, 
354989), 

Discussion. —Endocarps of this species differ from 
those of Palaeophytocrene by having much longer pro- 


trusions into the locule, and by the thinner endocarp 
wall (Pl. 19, figs. 12, 13). There is a very close resem- 
blance to endocarps of Pyrenacantha zenkeri (Sp. 
Moore) Exell from Angola, Africa, although the few 
specimens observed (e.g., Pl. 19, fig. 15) were smaller 
(only 15 mm long). Both are similar in having a thin 
endocarp with long protrusions into the locule. The 
genus Pyrenacantha has also been recognized in the 
Philippines (Sleumer, 1969). This is the first recogni- 
tion of Pyrenacantha from the North American fossil 
record. 


Family JUGLANDACEAE A. Rich ex Kunth. 


The Juglandaceae (walnut family) are a family of 
temperate to tropical trees with eight extant genera 
placed in four tribes: the Juglandeae (Juglans, Ptero- 
carya, Cyclocarya), Hicorieae (Carya), Engelhardieae 
(Engelhardia, Oreomunnea, Alfaroa), and Platycar- 
yeae (Platycarya). The family has an excellent fossil 
record of extant and extinct genera in the Tertiary of 
North America, Europe and Asia (reviewed by Man- 
chester, 1987a, 1989b). It is represented in the Nut 
Beds by wood of Engelhardioxylon (Manchester, 1983) 
and Clarnoxylon (Manchester and Wheeler, 1993) and 
by fruits of Cruciptera, Juglans, cf. Palaeocarya and 
Paleoplatycarya?, described below. 


Tribe JUGLANDEAE 
Genus CRUCIPTERA Manchester 


Cruciptera simsonii (Brown) Manchester 
Plate 21, figures 10-17 


Tetrapteris simsoni Brown 1940, pp. 344-356, fig. 9. 
Cruciptera simsonii (Brown) Manchester 1991, p. 719, figs. 1-11, 
13-15. 


Description.—Samara with four bract-wings radiat- 
ing in a plane surrounding a globose nutlet; fruit width 
including wings 25.0-38.0, avg. 33.2 mm (SD=6.45, 
n=5); pedicel 0.4 mm thick, at least 5.0 mm long, 
aligned with the axis of symmetry; nutlet globose ex- 
cept for an apical stylar projection, rounded basally, 
smooth, length 3.4-4.5 mm, equatorial diameter 3.6— 
5.0, avg. 4.2 mm (SD-0.51, n=6), nutlet unilocular; 
nutshell 0.2-0.3 mm thick, composed of isodiametric 
sclereids 15-30 um in diameter; base of the locule 
divided into four lobes by thick primary and secondary 
septa; secondary septum oriented in the plane of carpel 
closure, extending about ' of the distance from base 
to apex of the fruit; primary septum complete except 
for an aperture in the apical % of the nutlet; two la- 
cunae, more or less circular in cross section, situated 
near the periphery of the primary septum; plane of 
dehiscence dividing the nutshell into two halves, aligned 
with the secondary septum; wings attached in a plane 


about the equator ofthe nutlet; wings narrow, elongate, 
spatulate to parallel-sided, bluntly rounded to pointed 
at the apices; wing length 11.0-17.0 mm, width 2.5- 
4.6, аур. 3.5 mm (50-0.73, n=8), venation subpar- 
allel, somewhat spreading distally, without cross veins, 
occasionally dichotomizing, 15 to 20 veins across at 
the widest point ofthe wing, veins terminating distally 
without forming loops. 

Specimens.—OMSI Pb776, Pb899, Pb1175, Pb1580, 
UF 9263, 9612, samaras; UF 8867, permineralized 
nutlet; USNM 354375, locule cast. Holotype: USNM 
42334, a compression specimen from Clarno West 
Branch Creek locality. 

Discussion.— Cruciptera simsonii occurs at several 
localities of the Clarno Formation, including West 
Branch Creek, White Cliffs, Red Gap, Alex Canyon, 
Gosner Road, Dry Hollow and John Day Gulch (Text- 
fig. 2), and is also represented by a specimen from the 
middle Eocene Cashman locality of the Puget Group, 
western Washington (USGS loc. 9731, listed as Te- 
trapteris simsoni in Wolfe, 1968). The above descrip- 
tion is based only upon the Nut Beds specimens; fea- 
tures of the four sepals and two styles have been 
described previously based upon specimens from other 
Clarno localities (Manchester, 1991). Samaras with in- 
tact wings (Pl. 21, figs. 10-13) are relatively common 
in the basal leaf layer of Face 3, and a few isolated 
silicified nutlets and locule casts (Pl. 21, figs. 15-17) 
were found elsewhere in the locality. The silicified nut- 
lets reveal internal structure and anatomy but do not 
show the wings. Conversely, the impression specimens 
preserve the morphology and attachment of wing and 
petiole, but do not show internal details of the nutlet. 
One important specimen, however, shows both the 
wings (Pl. 21, fig. 11) and a well preserved nutlet (Pl. 
21, fig. 14) with the same internal structure as the 
isolated silicified nutlets, thus linking impression and 
petrifaction specimens as the same species. 

Transverse sections of the nutlets reveal a well de- 
veloped basal secondary septum in addition to the pri- 
mary septum dividing the locule into four chambers 
(Pl. 21, figs. 14-16). Accordingly, the associated chal- 
cedony locule casts are basally four-lobed (Pl. 21, fig. 
17), and are readily distinguished from the other small 
juglandaceous locule casts that occur in the Nut Beds, 
1.е., the two-lobed locule casts associated with Paleo- 
platycarya (Pl. 22, figs. 5-8) and Palaeocarya (Pl. 22, 
figs. 15, 16). The morphology of the nutlets and locule 
casts of this species are clearly juglandaceous. The nut- 
shell and septa are composed of isodiametric sclereids, 
differentiating them clearly from those of the tribe En- 
gelhardieae which are composed of fibers. The orien- 
tation of the wings in a plane perpendicular to the 
nutlet axis is a feature shared with extant Cyclocarya, 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


and the extinct Paleocene genus Polyptera (Manches- 
ter, 1989b, 1991). 

Cruciptera is also known from the Early Eocene of 
Wyoming, the Oligocene of Oregon and from the Mid- 
dle Eocene of Dorset, England (“Abelia sp.", Chandler, 
1964) and Messel, Germany (Manchester, 1991), and 
appears to predate the appearance of Pterocarya fruits 
(Late Eocene/Early Oligocene) in the fossil record. The 
foliage and pollen of Cruciptera remain to be deter- 
mined. Juglandaceous leaflets from the Nut Beds that 
were described and illustrated (Manchester, 1987a, p. 
89, fig. 38) as “Juglans (?) sp." were collected in the 
same stratigraphic layer as fruits of Cruciptera and are 
a candidate for the type of foliage produced by the 
extinct genus. Although the irregular spacing and course 
of secondary and tertiary veins is similar to that of 
extant leaflets of Juglans sect. Rhysocaryon, the shape, 
size, and venation of the teeth more closely resemble 
Pterocarya. Pterocarya fruits are absent from the entire 
Clarno formation; thus the foliage probably corre- 
sponds either with Juglans or Cruciptera. 


Genus JUGLANS L. 


Juglans clarnensis Scott 
Plate 21, figures 1—9 


Juglans clarnensis Scott 1954, p. 70, pl. 15 figs. 1-14; Bones 1979, 
pl. 2, fig. 6-8; Manchester 1987a, p. 107, fig. 46A-M. 


Description.— Nut globose to ellipsoidal, circular in 
cross section, unilocular, single-seeded, base rounded 
with slight depression in the attachment area, extreme 
apex sometimes with a small point; length 13.4-27.1, 
avg. 21.6 mm (SD-2.77, n=61), equatorial diameter 
15.4-23.4, avg. 19.3 mm (SD=2.01, n=65); surface 
with eight to 12 shallow longitudinal grooves joining 
at base and apex, sometimes anastomosing, including 
a pair of grooves corresponding to the plane of dehis- 
cence, nutshell 2-4 mm thick; primary septum per- 
pendicular to the plane of dehiscence with an aperture 
in the apical half, containing a pair of small longitu- 
dinal lacunae; secondary septum absent to rudimen- 
tary at the base of the locule in the plane of dehiscence; 
each valve of the nutshell containing a pair of laterally 
flattened longitudinal lacunae 4-8 mm wide and 1-1.5 
mm thick that join together at the base and apex; vas- 
cular supply of the primary septum axial in course. 
Locule cast smooth, quadrilaterally symmetrical about 
vertical axis, rounded basally, divided by the primary 
septum into two major lobes at the base, which are 
sometimes slightly emarginate basally by the faint de- 
velopment of a secondary septum; apex sharply point- 
ed, shoulder region concave to seldom convex. 


? See also Notes added in proof, p. 200. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 59 


Specimens.—Syntypes: UM 29906, 29907, 29908 
(Scott, 1954). Hypotypes: UF 4781, 4872, 4873, 4875, 
5338, 7965, 9424-9448, USNM 312378 (Bones, 1979, 
pl. 2), 354282-354291, 354293-354299, 354300- 
354315, 354361, 354362, nuts; OMSI Pb186, 188, 
189А, 237, 241, 251, UF 4874, 5339, 5340, 7965, 
9437, 9440, 9451-9469, UM 29908, USNM 354245- 
354246, locule casts. 

Discussion.— Juglans clarnensis is the oldest con- 
firmed fruit record of Juglans (Scott, 1954; Manches- 
ter, 1987a). There are more than 200 specimens ofthis 
Species in museum collections, preserved as nut casts 
(РІ. 21, figs. 1, 2) and molds, occasional permineralized 
huts (Pl. 21, figs. 3, 4), and locule casts (Pl. 21, figs. 5- 
9). Many more specimens ofthe esteemed Clarno wal- 
Nut reside in private collections. The specimens are 
most common in the middle and upper levels of Faces 
l and 2. Nuts that may represent the same species are 
Preserved as silica casts at the Brummers Spring lo- 
Cality (Text-fig. 2; Manchester, 1990). 

Criteria for placing this species in the extant genus 
Juglans were presented by Scott (1954) and Manches- 
ter (1987a). Juglans clarnensis possesses nutshell mor- 
Phology most similar to that of section Rhysocaryon 
(black walnuts) but has a less convoluted seed mor- 
Phology similar to that of to section Cardiocaryon (but- 
ternuts). 


Tribe ENGELHARDIEAE 
Genus PALAEOCARYA Saporta 


cf. Palaeocarya clarnensis Manchester 
Plate 22, figures 9-17 


Description.—Nutlet subglobose to obovoid, uniloc- 
ular, single-seeded, circular in cross section, base acute- 
Pointed, apex obtuse-pointed; smooth, except for a 
Single longitudinal rib preserved in a few specimens; 
fruit with a single incomplete septum dividing the base 
Of the locule and seed into two rounded lobes; primary 
Septum 0.4-0.6 mm thick, without lacunae, two vas- 
Cular bundles running axially through the septum ad- 
Jacent to the central axis; secondary septum absent; 
Nutshell thin (0.25 mm), without lacunae, composed 
of longitudinally oriented fibers; plane of dehiscence 
at right angles to the primary septum. Locule cast 
Smooth, quadrilaterally symmetrical about vertical axis, 
basally bilobed, each lobe corresponding to a cotyle- 
don, rounded basally and laterally, pointed apically, 
length 1.5-3.8, avg. 2.3 mm (SD=0.57, n=39), width 
Parallel to primary septum 1.7-3.4, avg. 2.6 mm 
(SD=0.48, n=40), width perpendicular to primary sep- 
tum 1.8-3.4, аур. 2.5 mm (SD=0.45, п=38). 

Specimens.—UF 9361, USNM 424640, 458416, 
More or less complete nutlets; HU 59994-59996, 


UCMP 10618-10621, UF 6456, 8747-8762, USNM 
354364 (4 specimens in matrix), 354366 (15 speci- 
mens), 354367, 354368 (2 specimens), 354369 (5 spec- 
imens), 354370 (11 specimens), 354371, 354372 (7 
specimens), 354374 (5 specimens), 354377 (3 speci- 
mens), 354378, 424640, 424703, locule casts. 

Discussion. — This species is known from numerous 
chalcedony locule casts (Pl. 22, figs. 9, 15, 16) and only 
a few partially complete permineralized nuts (Pl. 22, 
figs. 10-12, 14) which in section show the thin, smooth 
wall of the nutshell (Pl. 22, figs. 13, 17). Specimens 
were found in the upper part of Faces 1 and 3 of the 
Nut Beds, but not in the lower levels of finer grained 
sediments where samaras with intact wings would be 
likely to preserve. The juglandaceous morphology of 
the locule casts is clear from a simple comparison with 
the larger locule casts of Juglans clarnensis (Pl. 21, figs. 
5-9). The locule casts range greatly in size such that 
the smallest are about half the size of the largest, but 
no morphological differences were detected. Although 
none of the specimens have been found with remains 
of wings preserved, the small size is an indication that 
they are probably from winged fruits. The single rib 
remaining on some nutlets (Pl. 22, figs. 11, 14) may 
be a remnant of the midvein of a wing like that of 
Engelhardia. 

These nutlets and locule casts probably represent the 
engelhardioid taxon, Palaeocarya clarnensis, a winged 
fruit common at other localities of the Clarno For- 
mation (Manchester, 1987a). Without diagnostic char- 
acters from the wings, however, the Nut Beds material 
cannot be identified to Palaeocarya with certainty. The 
following lines of evidence, however, support the hy- 
pothesis that these fossils represent P. clarnensis: 1) We 
know that these were not the locule casts of Cruciptera, 
because Cruciptera fruits with intact nutlets show a 
basally four-lobed, not two-lobed, locule (cf. Pl. 21, 
fig. 17 with Pl. 22, fig. 9); 2) Affinities with Platycarya 
are also eliminated on the basis of locule cast mor- 
phology (cf. Pl. 22, figs. 5-8); 3) The nutlets of com- 
pressed fruits of P. clarnensis are highly variable in 
size, as are the isolated nutlets described above; 4) The 
nutshell and septa are composed of fibers, not sclereids, 
indicating affinities with the tribe Engelhardieae; 5) 
Wood of the Engelhardieae, the same tribe to which 
Palaeocarya belongs, is present in the Nut Beds (Man- 
chester, 1983). 

Palaeocarya is a generic name accommodating fossil 
fruits with characters found today both in the Asian 
genus Engelhardia and in the central American genus 
Oreomunnea. The fruits of these genera are samaroid, 
with a large trilobed wing developed from the inflo- 
rescence bract. If the species described above actually 
represents Palaeocarya clarnensis, then it appears that 


it was produced by a plant more similar to Engelhardia 
than to Oreomunnea. The bilobed locule casts of this 
species indicate a two-chambered nutlet like that of 
Engelhardia rather than an eight-chambered nutlet like 
that of Oreomunnea. 


Tribe PLATYCARYEAE 
Genus PALEOPLATYCARYA Manchester? 


Paleoplatycarya? hickeyi sp. nov. 
Plate 22, figures 1—8 


Etymology.—This species is named for Leo J. Hick- 
ey in recognition of his contributions to angiosperm 
paleobotany including the fossil record of Platycaryeae. 

Description.—Infructescence elongate, estimated 
length 26 mm, width 4.1-4.6 mm, composed of heli- 
cally arranged woody bracts on a relatively stout axis; 
one fruit with bilobed locule cast situated on the axil 
of each bract; bracts woody, shallowly v-shaped in 
cross section, ca. four to six bracts per cycle. Locule 
cast quadrilaterally symmetrical about vertical axis, 
obcordate in face view, pointed apically, bilobed ba- 
sally, laterally compressed in the plane opposite to the 
primary septum, length 1.6-1.8 mm, width 2.2 mm, 
thickness 0.7-1.0 mm, secondary septum not devel- 
oped, surface faintly verrucate. 

Specimens.— Holotype: UF 6454, infructescence with 
intact locule casts. Paratypes: UF 9251, USNM 353995, 
infructescence; UF 8763, USNM 354376, 435117, loc- 
ule casts. 

Discussion.— This species is represented by two cone- 
like infructescences (Pl. 22, figs. 1-3) and a few isolated 
locule casts (e.g., Pl. 22, fig. 6). The juglandaceous af- 
finity of the infructescences was not obvious until one 
of them was fractured to reveal the distinctive locule 
casts (Pl. 22, figs. 4, 5, 7, 8). The locule casts have the 
characteristic morphology of Juglandaceae, but are un- 
usual in being strongly compressed in the plane per- 
pendicular to the primary septum. This lateral flatten- 
ing may have been adaptive in the context of a wind- 
dispersed fruit. 

This species clearly belongs to the juglandaceous tribe 
Platycaryeae because of the cone-like infructescence 
with primary bracts persisting on the inflorescence axis. 
In all other tribes of the family, the primary inflores- 
cence bract becomes part of the fruit and does not 
persist on the infructescence axis. Although the Pla- 
tycaryeae have only a single modern genus, Platycarya 
of eastern Asia, the tribe was more diverse and wide- 
spread in the Tertiary, as documented by fruits, foliage 
and pollen from Europe and North America (Wing and 
Hickey, 1984; Manchester, 1987a, 1989b). The shal- 
low v-shape of the bracts in cross section (Pl. 22, fig. 
1) resembles that seen in the Eocene Platycarya ri- 
chardsonii (Bowerbank) Chandler (cf., Manchester, 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


1987a, fig. ТОН, I). The Nut Beds infructescence differs, 
however, from modern and fossil Platycarya species 
by its elongate, narrow construction. 

Similarly narrow, elongate infructescences with short 
bracts characterize Paleoplatycarya wingii from the 
early Eocene of Wyoming (Manchester, 1987a; Platy- 
carya castaneopsis sensu Wing and Hickey, 1984). 
However, features of the fruit wings and perianth con- 
sidered diagnostic of Paleoplatycarya (Manchester, 
1987a) are not preserved in the Nut Beds material, and 
no isolated fruits of Paleoplatycarya are known from 
the Clarno Formation. Pending the recovery of more 
informative specimens, it seems appropriate to assign 
this species to **Paleoplatycarya?". 

Among extensive collections from other Clarno lo- 
calities, I have observed only two other infructescences 
that appear to represent P.? hickeyi. Both are impres- 
sion specimens from West Branch Creek (UF 9871, 
9872). Another type of infructescence that is relatively 
common at West Branch Creek, named Platycarya 
manchesteri by Wing and Hickey (1984), is superfi- 
cially similar to that described above. However, anal- 
ysis of numerous specimens of P. manchesteri has 
shown that each nutlet is surrounded by four bract- 
like structures, rather than just one, indicating that the 
affinities are not with Platycarya. The arrangement of 
the bract-like structures suggests that they may be im- 
mature infructescences of Cruciptera. 

It is possible that P?. hickeyi infructescences pro- 
duced fruits of the extinct platycaryoid genus Hooleya. 
Although the distinctive biwinged fruits of Hooleya 
lata Wing and Hickey (1984) occur at the West Branch 
Creek and White Cliffs localities of the Clarno For- 
mation (Text-fig. 2), they have not been identified from 
the Nut Beds. Locule casts of P.? hickeyi are compa- 
rable in size and shape, however, to the nutlet body 
seen in the compressed Hooleya fruits (cf., Manchester, 
1987, figs. 12А-Е, С, Н). Clarnoxylon, an extinct type 
of juglandaceous wood known both from the Nut Beds 
and from West Branch Creek, possesses crystaliferous 
idioblasts characteristic of Platycaryeae (Manchester 
and Wheeler, 1993), and is a likely candidate for the 
type of xylem produced by Hooleya and/or Paleopla- 
tycarya trees. 


Family LAURACEAE Juss. 


The Lauraceae are a family of about 45 genera and 
2200 species concentrated in the tropics and warm 
temperate areas. The family is first reliably reported 
from the Cretaceous (Drinnan et al., 1990; Herendeen, 
1991) and was widespread by the early Tertiary. The 
Lauraceae are represented in the Nut Beds by at least 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 61 


two types of foliage, and wood of Ulminium (Scott and 
Wheeler, 1982), as well as by the fruits described be- 
low. 

Fruits of Lauraceae are unilocular and single-seeded 
and the seed is exalbuminous with a pair of large hemi- 
Spherical cotyledons. Reid and Chandler (1933) pro- 
vide an excellent synopsis of the fruit and seed char- 
acters of extant Lauraceae. The fruit may or may not 
be enclosed partially or completely in the hypanthium 
or “cupule.” A prominent scar may remain at the base 
of the epicarp where it detaches from such a cupule. 

he endocarp typically has a uniseriate layer of anti- 
Clinally oriented columnar cells. This layer, which may 
be thick, thin or intermittent, generally appears to form 
the innermost coat of the pericarp. Oil cells, a feature 
ОЁ xylem and other tissues of Lauraceae, may also be 
Present in the cupule and pericarp. The parenchyma 
От sclerenchyma cells are often oriented radially around 
oil cells, forming the “‘stellate clusters of cells” referred 
to by Reid and Chandler (1933) as a diagnostic feature 
ОЁ the family. 

Although it is fairly easy to identify fruits to the 
Lauraceae, this family is notoriously difficult when it 
Comes to making generic assignments. Part of the dif- 
ficulty, which applies even to extant species, is that 
“Asa rule, both floral and fruit characters are necessary 
for identification, but a single specimen rarely has both 
flowers and fruits” (van der Werff, 1991). Therefore, 
With the exception of Lindera, the Nut Beds species 
have been assigned to the stereotypic genera Lauro- 
calyx and Laurocarpum. 


Genus LINDERA Thunb. 


Lindera clarnensis sp. nov. 
Plate 23, figures 1-8 


Etymology. — The epithet indicates the source of the 
Species, the Clarno Formation. 

Description. —Endocarp spheroidal, bilaterally sym- 
Metrical, with a faint ridge running from apex half way 
to base in plane of symmetry; unilocular, single-seeded; 
length 3.8-6.0, avg. 5.2 mm (SD=0.83, п=5), equa- 
torial diameter 3.8-6.3, avg. 5.3 mm (SD=0.98, n=5); 
Endocarp thin-walled, composed of a uniseriate layer 
of Columnar cells, these cells about 100 um high and 
About 15 um wide. Seed exalbuminous, filled com- 
Pletely by a pair of large hemispherical cotyledons. 

Specimens.—Holotype: USNM 435006. Paratypes: 

SNM 335472, 435007, 435008, 424728. 

Discussion.—The combination of unilocular, glo- 

Ose endocarps (Pl. 23, figs. 1-4) with a prominent 
columnar layer (Pl. 23, figs. 6-8), a single seed and 
arge plano-convex cotyledons (Pl. 23, fig. 5) confirms 
that this species belongs to the Lauraceae. Most extant 


species of Lauraceae that were examined for this study 
are axially elongate, rather than globose. Some extant 
species of Cinnamomum and Lindera, however, have 
small globose endocarps comparable in size and in- 
ternal structure to the fossil. The endocarp wall of Cin- 
namomum camphora (L.) Presl is much thicker than 
that in Lindera spp. This fossil is virtually indistin- 
guishable from fruits of Lindera umbellata Thunb. from 
Japan and L. obtusiloba Blume from the Philippines. 
Because of the high degree of similarity to these species, 
and since I have not found precisely similar species in 
a survey of other extant genera, the Nut Beds species 
is assigned to the extant genus. 


Genus LAUROCALYX Reid et Chandler 


This genus was established by Reid and Chandler 
(1933) for “cupuliferous fruits belonging to the family 
Lauraceae which, for one cause or another, cannot be 
referred to living genera.” The preservation of a per- 
sistent perianth or cupule distinguishes this fossil genus 
from Laurocarpum. 


Laurocalyx wheelerae sp. nov. 
Plate 23, figures 9-15 


Etymology.—This species is named for Elisabeth A. 
Wheeler, recognizing her contributions to the analysis 
of woods from the Clarno Formation, including those 
of the Lauraceae. 

Description.—Fruit prolate, circular in transverse 
section, rounded apically and basally, unilocular, sin- 
gle-seeded, length 7.7-10.1 mm, width 7.9-10.9 mm, 
thickness 7.4-10.9 mm; embedded in a thick cupule 
covering the basal Ys of the fruit; endocarp surface 
smooth, with a prominent, recessed, elliptical to ren- 
iform basal scar approximately 4.5 mm wide and 2.8 
mm thick containing three vascular bundles arranged 
in a triangular pattern, endocarp wall about 0.7 mm 
thick, and composed ofa uniseriate layer of anticlinally 
elongate cells; inside of seed with solid core of rounded 
parenchyma cells, 38-50 um in diameter, this core is 
surrounded by larger angular parenchyma cells, to the 
outside of this is a uni- to biseriate seed coat of dark- 
pigmented periclinally elongate rectangular cells 20- 
22 um high, 20-38 um wide. Cupule composed mostly 
of isodiametric parenchyma cells 38-50 um in diam- 
eter with interspersed larger cells 60-80 um in diameter 
with black contents interpreted as oil cells. 

Specimens.—Holotype: UF 9406. Paratypes: UF 
9407-9410, USNM 355626, 355630, 355631, 355634, 
355638, 435085, 424807. 

Discussion.— Laurocalyx wheelerae is represented by 
permineralized fruits and locule casts, sometimes with 
the cupule attached. Isolated endocarps and locule casts 
are recognizable even without the attached cupule be- 
cause of the conspicuous elliptical attachment scar (Pl. 


62 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


23, figs. 9, 11). The holotype is relatively complete, 
including both the endocarp and the cupule. The en- 
docarp was removed from the cupule for comparison 
with other isolated endocarps (Pl. 23, figs. 11, 12), and 
was subsequently reassembled with its cupule and sec- 
tioned longitudinally, showing the lateral extent of cu- 
pule coverage (Pl. 23, fig. 13), the presence of oil cells 
within the cupule (Pl. 23, fig. 14) and the layer of 
columnar cells in Ше endocarp (Pl. 23, fig. 15). These 
features confirm that the species represents Lauraceae. 


Genus LAUROCARPUM Reid et Chandler 


Laurocarpum Reid et Chandler is a stereotype genus 
erected to accommodate unilocular, single-seeded fruits 
that have characters diagnostic of Lauraceae, but that 
cannot be placed in a particular modern genus (Reid 
and Chandler, 1933). At least three species from the 
Nut Beds flora are assignable to this genus: L. han- 
cockii, L. nutbedensis, and L. raisinoides, which are 
described below. 

Scott (1954) identified one specimen from the Nut 
Beds to Laurocarpum proteum Reid et Chandler, a 
species based on well preserved specimens from the 
London Clay flora (Reid and Chandler, 1933). The 
specimen is an ovoid locule cast 11 mm long and 9 
mm in diameter with some fragments of endocarp re- 
maining. The shape of the apex and the truncate base 
with a large circular chalaza compare favorably with 
the London Clay species. I have reexamined the spec- 
imen figured by Scott (1954). Although external mor- 
phological features of the locule cast seem to support 
assignment to Lauraceae, it is unfortunate that char- 
acters of the cotyledons and wall layers remain un- 
available to confirm the assignment. I have been un- 
able to locate any additional specimens that might 
provide more characters upon which to confirm the 
systematic position of this fossil. 


Laurocarpum hancockii sp. nov. 
Plate 24, figures 1—3 


Etymology.—The epithet is in memory of Alonzo 
W. Hancock, who established a field station promoting 
paleontological field work in the Clarno Formation. 

Description.— Fruit pyriform, circular in cross sec- 
tion, unilocular, single-seeded, length 25 mm, equa- 
torial diameter 18.5 mm; fruit wall 2.0 mm thick, com- 
posed of two layers: an inner uniseriate layer of 
anticlinally oriented columnar cells, 250 um high and 
62-112 um wide, and an outer layer, presumably the 
mesocarp (exocarp not preserved), composed of a mass 
of thin-walled parenchyma cells 20-82 um in diameter, 
arranged in radiating patterns around solitary or clus- 
tered stone cells, the stone cells or isodiametric scler- 
eids 50-100 um in diameter, and occasional oil cells 


with dark contents; seed filled by two large hemispher- 
ical cotyledons. Cupule not preserved. 

Specimen.— Holotype: OMSI Pb262. 

Discussion.— Laurocarpum hancockii, known from 
a single specimen, has the largest fruit of the lauraceous 
taxa observed from the Nut Beds. The fruit is clearly 
lauraceous because of the two large hemispherical cot- 
yledons (Pl. 24, figs. 1, 2), the layer of columnar cells 
lining the locule (Pl. 24, fig. 3), and the presence of 
stone cells about which the surrounding parenchyma 
is stellately arranged (Pl. 24, fig. 3). 


Laurocarpum nutbedensis sp. nov. 
Plate 24, figures 4—7 


Etymology.—The epithet nutbedensis refers to the 
Nut Beds locality. 

Description.—Fruit prolate, unilocular, single seed- 
ed; length 8.0 mm, equatorial diameter 6.8 mm; sur- 
face relatively smooth; pericarp ca. 1.0 mm thick, con- 
sisting of: a thin uniseriate outermost layer of 
anticlinally oriented columnar cells 40-50 um high, 
10-15 um wide, a main layer 750-950 um thick com- 
posed of isodiametric parenchyma cells, typically 20- 
50 um in diameter, but smaller and tangentially flat- 
tened (16-18 um high) toward the inside of this layer, 
large oil cells 50-70 um in diameter interspersed among 
this tissue; this layer lined on the inside by a uniseriate 
layer of columnar cells, and then in turn by a layer of 
several rows thick of rectangular cells, 200 um thick; 
exocarp not preserved. Two hemispherical cotyledons 
filling the seed cavity. 

Specimens.— Holotype: USNM 435090. 

Discussion.— Laurocarpum nutbedensis is repre- 
sented by a single permineralized specimen that has 
been sectioned transversely (Pl. 24, figs. 4-7). Laura- 
ceous affinity is suggested by the columnar layers, oil 
cells, and single seed with hemispherical cotyledons. 
Oil cells of the pericarp are slightly larger and more 
rounded than the surrounding parenchyma cells. 


Laurocarpum raisinoides sp. nov. 
Plate 24, figures 8-15 


Etymology.—The epithet refers to the wrinkled, rai- 
sin-like surface of the fruit. 

Description. — Fruit subglobose, rounded apically and 
basally, unilocular, single-seeded, length 4.5 mm, 
equatorial diameter 4.0-5.2 mm; surface longitudi- 
nally wrinkled; pericarp with an inner uniseriate layer 
of columnar cells 20 um thick giving rise to a 4-5- 
seriate layer of small thick walled isodiametric cells 
7.5-10 um, followed by a mesocarp layer of paren” 
chymatous, cells 17-25 um in diameter, with numer- 
ous interspersed oil cells 37-58 um in diameter; ехо“ 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 63 


Carp uniseriate, composed of of anticlinally oriented 
columnar cells 21 um high, 5 um wide. 

Specimens.—Holotype: UF 5672. 

Discussion.—Laurocarpum raisinoides is represent- 
ed by one very well preserved specimen with clear 
affinities to the Lauraceae. The longitudinal wrinkles 
Оп the outer surface (Pl. 24, figs. 8-10) suggest a leath- 
ry pericarp. Sections through the mesocarp reveal 
abundant oil cells that are larger than the surrounding 
Parenchyma cells and filled with dark contents (Pl. 24, 
figs. 12-1 5). The endocarp and exocarp layers are com- 
Posed of columnar cells (Pl. 24, figs. 13, 14). Fruits of 
extant Cinnamomum camphora are morphologically 
and anatomically similar; it is likely, however, that 
Similar fruits could be found among various different 
extant genera. 


Family LEGUMINOSAE Juss. 


The Leguminosae are represented only by one type 
Of fruit from the Nut Beds locality. Three additional 
legume fruit types are known from the Clarno For- 
mation (West Branch Creek and Gosner Road locali- 
tes), and remain to be described. Wood similar to 
Acacia from the Nut Beds (Wheeler and Baas, 1992, 

8. 9-13) indicates that arborescent legume taxa were 
Present, The low diversity and relatively low number 
Of specimens of legume fossils in the Clarno Formation 
Suggest that legumes were a relatively minor compo- 
Dent of the flora. This is in contrast to the diversity of 
“gume fruits and foliage known from sediments of 
Similar age in southeastern North America (Herendeen 
and Dilcher, 1990). 


Genus LEGUMINOCARPON Goeppert 


Leguminocarpon sp. 
Plate 25, figure 9 


Description. — Fruit elliptical, bilaterally symmetri- 
Cal, laterally flattened in plane of symmetry, about 35 
Mm long, 12 mm wide, and a few mm thick, base and 
apex rounded; fruit narrowed over the basal 3, possibly 
Where some ovules aborted; fruit body with very fine 
Transverse veins. 

Specimen.—OMSI Pb1725. 

Discussion.—This specimen is an impression in silt- 
Stone from in the leaf layer at the base of Face 3. The 
texture indicates that it may have been thick-textured, 
Possibly leathery, but not woody. Although it is clearly 
а pod of the Leguminosae, it does not show the position 
От orientation of seeds, and on the basis of this single 
Specimen it is not possible to determine the intrafam- 
Ша] relationships. Lacking additional characters, it is 
Unlikely that the intrafamilial affinities can be deter- 
Mined. Bones (1979, pl. 3, fig. 4) figured another pos- 


sible legume pod from the Nut Beds (USNM 326716), 
but the preservation is not good enough to confirm its 
affinities. 


Family LYTHRACEAE Jaume St-Hil. 


The Lythraceae are a family of about 26 genera of 
herbs, shrubs and trees with temperate and tropical 
distribution around the world. The family has an ex- 
cellent record in the Tertiary of Europe and North 
America. Both extant and extinct genera have been 
recognized on the basis of isolated seeds, as reviewed 
by Tiffney (1981), Friis (1985) and Cevallos-Ferriz and 
Stockey (1988a). In the Nut Beds this family is rep- 
resented by fruits and seeds of Decodon. In addition, 
there is some evidence that seeds of Tripartisemen (p. 
113), which are very common at the Nut Beds, belong 
to the Lythraceae. 


Genus DECODON Gmelin 


Decodon sp. 
Plate 25, figures 1-8 


Description.—Fruit a thin-walled, globose, multi- 
seeded capsule, 2.3-3.9, avg. 3.4 mm (SD=0.41, n=5) 
in diameter, with hypogynous perianth; pedicel at least 
3 mm long and 0.4 mm in diameter; fruit multilocular, 
the number of locules undetermined, with thin septa 
(0.8-1.2 mm) radiating from a thickened central axis 
in the basal % of fruit, placentation axile, seeds nu- 
merous; fruit wall thin, 0.8-1.2 mm, approximately 
equal in thickness to the septa. Seeds anatropous, an- 
gular in section, laterally compressed, rounded dor- 
sally, often narrowed and forming a keel ventrally, 
length 0.7-1.1 mm, width 0.4-0.6 mm; seed coat com- 
posed of three layers: 1) innermost layer lining the 
embryo cavity 18 um thick, composed of a uniseriate 
layer of cuboidal cells; 2) middle layer, forming the 
body of the seed through which the raphe travels, mul- 
tiseriate, composed of isodiametric cells 10-25 um in 
diameter; 3) outer layer uniseriate, composed of thin- 
walled rectangular cells, this layer 10 шт thick over 
most of the seed, but dramatically thickened up to 120 
um to form the ventral germination valve, cells of the 
germination valve anticlinally elongate, 70-120 um 
long and 30 um wide; embryo cavity curved, nearly 
circular or elliptical in cross section but with a straight 
ventral margin adjacent to the germination valve; ra- 
phe passing through the middle layer of the seed. 

Specimens. — UF 6449, pedicellate fruit; UF 5668, 
6450, 6451, 9287, fruits with intact seeds. 

Discussion.—Several permineralized fruits of De- 
codon have been recovered from the Nut Beds. Some- 
times the seed tissues are not preserved and embryo 
casts are preserved within the empty or carbonaceous 


64 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


mold of the seed (Pl. 25, fig. 1). Serial sections, how- 
ever, provide excellent details of fruit and seed anat- 
omy (Pl. 25, figs. 2, 4-8). One of the specimens was 
sectioned longitudinally and clearly shows the pedicel, 
hypogynous calyx, a thin septum and the thin fruit wall 
(Pl. 25, fig. 5). These features of the fruit, together with 
axile placentation and small anatropous seeds with 
ventral germination valves and curved embryo cham- 
bers, indicate affinties with the Lythraceae. 

The structure and anatomy of the fossil compare 
favorably with extant Decodon verticillatus (L.) Ell. (cf., 
Friis, 1985, pl. 15, figs. 10-12; Cevallos-Ferriz and 
Stockey, 1988a, figs. 19, 20). The Clarno species is 
distinguished from permineralized remains of D. al- 
lenbyensis Cevallos-Ferriz and Stockey, from the Eo- 
cene of Princeton, southern British Columbia, in the 
shape of the embryo cavity. In the Clarno fossil, as in 
D. verticillatus, the embryo cavity usually is D-shaped 
in transverse section because of the relatively straight 
inner face of the germination valve, but in D. allen- 
byensis the embryo cavity typically is O-shaped in 
transverse section, evidently reflecting a convex inner 
margin of the germination valve. 

Decodon has a single extant species, D. verticillatus, 
that is an aquatic shrub of swamps and marshy areas 
distributed in eastern North America from Ontario to 
central Florida. The genus was formerly more widely 
distributed, with many seed records in the Tertiary of 
Europe (Mai and Walther, 1978; Friis, 1985). 


Family MAGNOLIACEAE Juss. 


The Magnoliaceae are a family of seven genera and 
about 200 species of trees and shrubs distributed from 
the Himalayas to Japan and western Malesia, eastern 
North America to tropical America. The family is di- 
vided into two subfamilies (Nooteboom, 1985): the 
Magnolieae (Elmerrillia, Kmeria, Magnolia, Manglie- 
tia, Michelia, Pachylarnax) and the Liriodendreae 
(Liriodendron). Other previously proposed genera of 
Magnolieae, such as Talauma (— Magnolia), are dis- 
missed by Nooteboom (1985). The Magnoliaceae are 
represented in the Nut Beds by at least three species 
of Magnolia. Although wood of Liriodendron has been 
reported from the Clarno “Osmunda” locality (Scott 
and Wheeler, 1982; locality data from Wheeler, pers. 
comm., 1977), no fruits or seeds of the genus have 
been recovered from the Nut Beds or any other Clarno 
localities. 


Genus MAGNOLIA L. 


Leaves of Magnolia are reported from several other 
Tertiary localities in western North America (Taylor, 
1990), but, unless confirmed by cuticular characters, 
the identification of leaves to Magnolia should be 


viewed with caution because Magnolia leaves are en- 
tire-margined and have a brochidodromous pattern of 
venation that occurs in many dicotyledonous families. 

Two species of Magnolia have been identified from 
the Nut Beds on the basis of woods: Magnolia longi- 
radiata and M. angulata (Scott and Wheeler, 1982). 
The xylem of both species appears primitive in relation 
to that of extant Magnoliaceae in having more bars per 
scalariform vessel perforation (Scott and Wheeler, 
1982). 

I recognize three species of Magnolia on the basis 
of seeds from the Nut Beds. There is considerable vari- 
ation in shape and size among them, however, and it 
is likely that more species could be recognized using 
the fine-splitting criteria previously applied for seeds 
from the London Clay flora (Reid and Chandler, 1933; 
Chandler, 1961b; see Chandler, 1978, pp. 7—9 for her 
philosophy of taxonomic splitting as applied to Mag- 
nolia). The terminology for describing Magnolia seeds 
used below follows that of Tiffney (1977). 


Magnolia muldoonae sp. nov. 
Plate 26, figures 1-9, 18, 20 


Etymology.—This species is named for Maureen 
Muldoon, recognizing the importance of her assistance 
with field work at the Nut Beds. 

Description.—Infructescence an elongate axis with 
helically arranged woody follicles; seeds narrowly to 
broadly ovate, asymmetrical, often dorsiventrally 
compressed, lensoidal to rounded-triangular in cross 
section, acutely to obtusely pointed at the micropylar 
end, rounded basally with a slight bulge at the heter- 
opyle; length 4.0-7.8, avg. 6.0 mm (SD=0.93, п-58), 
width 2.5-8.4, avg. 4.7 mm (SD=1.44, п=61), thick- 
ness 1.5-3.7, avg. 2.4 mm (SD=0.49 n=60); seed cast 
surface smooth, heteropyle located centrally on the 
rounded end with a relatively small, circular moat, 
stalk bowed ventrally at right angles to the long axis 
of seed; sclerotesta 250 um thick. 

Specimens.— Holotype: UF 6529, infructescence with 
intact seeds. Paratypes: UF 6546, 6555, 6556, 9728 
(22 specimens), USNM 312756 (Bones, 1979, pl. 5; 
fig. 5), 355378 (6 specimens), 355407 (3 specimens). 
355413 (2 specimens), 355421, 355537 (12 speci- 
mens), 355540 (20 specimens), 355646 (several spec 
imens), 424697-424699, seed casts. 

Discussion.— Magnolia muldoonae is the most сот" 
mon species of Magnolia in the Nut Beds assemblage: 
and is particularly abundant in a stratum exposed 10 
the gully between Faces 3 and 4. It is represented bY 
a permineralized infructescence (Pl. 26, figs. 1, 2) and 
numerous chalcedony seed casts (e.g., Pl. 26, figs. a 
9). The seed casts are usually internal molds of the 
sclerotesta, and thus present slightly different surface 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 65 


features from the lignitized fossils commonly described 
In the literature (Mai, 1975; Tiffney, 1977). Although 
the seeds vary from longer than wide (Pl. 26, figs. 3- 
5) to wider than long (Pl. 26, figs. 6-8), similarities of 
the heteropylar region, including the slightly swollen 
Chalaza and ventrally directed stalk (Pl. 26, figs. 7, 8), 
Suggest that they all represent the same species. This 
18 also suggested by the occurrence of numerous seeds 
of variable length/width ratio together in concentrated 
Pockets in the sediment. 

Tiffney (1977) presented a helpful conspectus of ex- 
fant and fossil magnoliaceous seeds. Seeds with a 
Smooth-surfaced sclerotesta characterize most extant 
Species of Magnolia. Seeds of other species of Mag- 
nolia, including those formerly placed in Talauma, 
ànd seeds of Michelia and Manglietia, have verrucate 
and/or rugulate surfaces. This surface ornamentation 
Carries through to the inside of the sclerotesta, and 
therefore the smooth surface of the M. muldoonae seed 
Casts supports the assignment to Magnolia. 

) The thickness of the sclerotesta as seen in thin-sec- 
tions of the infructescence is up to 250 um, as is typical 
for most extant species of Magnolia. Nevertheless, the 
Sclerotesta is thin in relation to that of some other fossil 
Species, such as M. waltonii Tiffney (1977) from the 
Oligocene Brandon Lignite of Vermont and М. lignita 
from the Miocene and Pliocene of Europe (Mai, 1975), 
Which may be 400 to 1000 um thick. Tiffney (1977, p. 
319) reported a single seed from the Clarno Nut Beds 
locality (HU 59970) that is similar to М. waltonii in 
Size, testal wall thickness and chalazal configuration, 
but noted that further comparison is limited by poor 
Preservation of the Clarno specimen. The specimen 
May represent an additional species of Magnolia that 
IS not considered in this monograph due to insufficient 
Material. 


Magnolia paroblonga sp. nov. 
Plate 26, figures 10-17 


Etymology.— Para (Gr = beside, near) + oblonga, 
denoting the similarity of this species to M. oblonga 
Chandler from the London Clay flora. 

Description.—Seed cast narrowly to widely subcor- 
diform, asymmetrical, dorsiventrally compressed, more 
Ог less lenticular in cross section; length 5.0-7.3, avg. 
6.3 mm (SD=0.63, n=17), width 5.5-11.3, avg. 8.5 
mm (SD=2.0, n=17), thickness 2.0-4.3, avg. 3.3 mm 
(SD=0.7 1, n=17); base truncate, with an elongate cha- 
lazal groove that is nearly as long as the seed width, 
With (in two specimens, UF 9190, 9192) a small central 
elliptical stalk scar; apex with acute to obtuse micro- 
Pylar point; lateral margins straight to rounded, some- 
times with one or two flat faces (suggesting more than 
Опе seed per carpel), often with a faint median ventral 


groove; inner surface of sclerotesta smooth or with 
longitudinal striations the chalazal scar. 

Specimens.— Holotype: UF 9192. Paratypes: UF 
9184-9191, 9192-9196, 9250, USNM 434998- 
435000. 

Discussion. — Magnolia paroblonga is represented in 
the Nut Beds assemblage by internal molds ofthe seed 
coat collected at the top of Face 3. The thickness of 
the sclerotesta is not known. It is readily distinguished 
from the other Clarno magnolias by longitudinal stri- 
ations in some of the specimens (Pl. 26, figs. 14-16) 
and by the elongate chalazal moat (Pl. 26, figs. 14, 15, 
17). This elongate groove is a feature shared with М. 
longissima (Bowerbank) Reid et Chandler (1933) and 
M. oblonga Chandler (196 1b) of the London Clay flora. 
Many of the Nut Beds specimens conform with Chan- 
dler’s diagnosis of M. oblonga, but there are also sev- 
eral specimens that are narrower and thicker than those 
described by Chandler. 

Seeds with an elongate chalazal moat are very un- 
usual among extant Magnolia species. In reviewing 
more than 30 species represented in the extant Mag- 
nolia seed collection prepared by Bruce Tiffney at the 
Harvard Herbaria, I observed only a single species with 
this character, Magnolia quinquepeta (Висћог) Dandy 
(Arnold Herbarium: Ren Chang Ching 2037) from 
Chekiang, China. The structure is not obvious on the 
surface of the sclerotesta, but is seen on the inside 
surface as a fusiform thickening at the heteropylar end 
of the endosperm cavity. 


Magnolia tiffneyi sp. nov. 
Plate 25, figures 10—12; 
Plate 26, figure 19 


Etymology.—This species is named in recognition 
of the contributions of Bruce H. Tiffney to the study 
of fossil Magnolia seeds. 

Description.—Seed narrowly to broadly ovate, 
asymmetrical, often dorsiventrally compressed, len- 
soidal to rounded-triangular in cross section, some- 
times with a median longitudinal trough on the ventral 
side, acutely to obtusely pointed at the micropylar end, 
rounded at the heteropylar end; length 8.4-12.1, avg. 
10.3 mm (SD=1.15, n=10), width 6.0-10.2, avg. 8.3 
mm (SD=1.44, n=10), thickness 2.6-4.5, avg. 3.5 mm 
(SD=0.71, n=8), surface smooth; heteropyle located 
centrally on the rounded end, not swollen, with a prom- 
inent v-shaped moat with stalk oriented more or less 
parallel to the long axis of seed. 

Specimens.— Holotype: UF 6549, seed cast. Para- 
types: UCMP 10744, UF 9729-9732, USNM 435002, 
435003, UWBM 36070, seed casts; UF 9733, infruc- 
tescence. 

Discussion.— Although M. muldoonae is relatively 


common in the Nut Beds, М. tiffneyi is represented 
only by a few specimens. Seeds of Magnolia tiffneyi 
differ from those of M. muldoonae by their large size, 
the v-shaped rather than rounded moat, and by the 
lack of a chalazal swelling. In addition, the stalk is 
oriented more or less parallel to the long axis of the 
seed rather than bending ventrally (Pl. 25, figs. 11, 12). 
Seeds of M. tiffneyi are similar in size and morphology 
to those of extant M. grandiflora L. of the eastern Unit- 
ed States, and to fossils of М. septentrionalis Tiffney 
from the Brandon Lignite of Vermont (Tiffney, 1977). 


Family MENISPERMACEAE Juss. 


The Menispermaceae, or moonseed family, includes 
about 78 living genera and 520 species. The family 
consists mostly of woody climbers and, with the ex- 
ception of a few temperate species, is chiefly tropical, 
with representatives in Asia, Africa, Australia and 
America. Menisperms are common “‘where the canopy 
is interrupted, e.g. along river banks, margins of forest 
or in clearings” (Forman, 1986). The Menispermaceae 
are diverse in the Eocene of Europe as indicated by 15 
genera and 22 species recognized in the London Clay 
(Chandler, 1964). Based upon leaves Wolfe (1977) rec- 
ognized 11 genera of Menispermaceae from the Paleo- 
gene of Alaska. 

Fruits of the family are drupes with distinctive en- 
docarps that provide characters important in distin- 
guishing tribes and genera (Diels, 1910; Thanikaimoni, 
1984; Forman, 1986). The endocarps are commonly 
boat-shaped or horseshoe-shaped with various pat- 
terns of ornamentation or sculpture and often have a 
prominent bilobed ventral infold or condyle. 

The Menispermaceae are the most diverse family in 
the Nut Beds with three tribes and 13 genera of fruits: 
Coscineae (Anamirta), Tinosporeae (Chandlera, 
Odontocaryoideae, Calycocarpum, Tinospora, Tinom- 
iscoidea, Atriaecarpum, Curvitinospora) and Menis- 
permeae (Davisicarpum, Diploclisia, Eohypserpa, Pa- 
leosinomenium, Thanikaimonia). Scott (1954, 1956) 
described two of the extinct genera, Chandlera, and 
Odontocaryoidea, and noted their importance in un- 
derstanding evolution of endocarp sculpture in the Ti- 
nosporeae. In addition, at least four types of menis- 
permaceous leaves are present (Manchester, 1981 and 
unpublished). 


Tribe COCSINIEAE 
Genus ANAMIRTA Colebr. 
Anamirta leiocarpa sp. nov. 
Plate 27, figures 1-7; Text-figure 14A 
Etymology.— Leio (Gr = bald) + karpos (Gr = fruit), 
emphasizing the smooth surface of the endocarp. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Description. — Fruit unilocular with a single seed; en- 
docarp globose, diameter 6.2-8.6, avg. 7.5 mm, 
(SD=0.6 mm, n=5), smooth except for a fine median 
keel, and the closely adjacent pedicel and style scars; 
locule subspherical, deeply intruded by a bilobed bul- 
bous condyle; condyle about % as high and У to Y. as 
wide as the endocarp and hollow from the base with 
a median septum, wall of the condyle 0.2-0.3 mm 
thick; endocarp wall 0.32-0.40 mm thick, composed 
of fibers 5.0-8.5 um in diameter, 250 um long that are 
oriented periclinally in the М adjacent to Ше locule, 
but anticlinally oriented in the outer 3⁄4 of the wall. 

Specimens.— Holotype: UF 8745. Paratypes: UF 
5712, 6305-6309, 6468, 8728, 8746, USNM 355622, 
424641, UCMP 10612, 10613. 

Discussion.—Anamirta leiocarpa is represented in 
the Nut Beds by silicified endocarps. Because the outer 
surface of these endocarps is smooth and rather non- 
descript (Pl. 27, fig. 1), the species is most easily rec- 
ognized in broken or sectioned specimens showing the 
prominent, central, bilobed condyle (Pl. 27, figs. 2-5). 
Thin sections reveal that the endocarp wall is com- 
posed predominantly of anticlinally oriented fibers (Pl. 
27, fig. 7). The identification of these fossils as Ana- 
mirta is supported by the globose form and relatively 
thin wall of the endocarp, the deeply intruded condyle 
and by the wall anatomy. Anamirta leiocarpa resem- 
bles the only modern species of the genus, A. cocculus 
(L.) Wight et Arn. (Pl. 27, figs. 8, 9), in size and en- 
docarp morphology, but A. cocculus has a minutely 
verrucate surface unlike the smooth surface of the fos- 
sil. 

Extant Anamirta is distributed from India to Indo- 
china and throughout Malesia. A. cocculus is an in- 
habitant of lowland areas that occurs, according to 
Forman (1986) “in a variety of conditions, on banks 
of rivers and streams, coastal forest, savanahs, on ba- 
salt, limestone and sandy soil, both in rain-forest con- 
ditions and in seasonal climates. . . , but with a distinct 
preference for seasonal conditions." Although this is 
the first confirmation of Anamirta based on the fossil 
fruit record, the genus has also been identified on the 
basis of fossil leaves from the Eocene Kulthieth For- 
mation of southeastern Alaska (Wolfe, 1977). 


Tribe TINOSPOREAE 
Genus CHANDLERA Scott 


Chandlera lacunosa Scott 
Plate 27, figures 10—20; Text-figure 13B 


Chandlera lacunosa Scott 1954, pp. 76-78, pl. 15, figs. 32-41, Scott, 
1956, pp. 75-77, fig. 1а. 


Description.— Fruit unilocular with a single seed; en- 
docarp oblong, bilaterally symmetrical, rounded-rect- 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 67 


angular in dorsiventral view, long-elliptical in lateral 
view, elliptical in cross section, rounded apically and 
basally; length 26.5-51.7, avg. 32.7 mm, (SD=4.75, 
n=22), width 14.7-28.9, avg. 24.3 mm (SD=4.5, n=27), 
thickness 9.8-23.5, avg. 15.9 mm (SD=3.1, n=27); 
Surface smooth, with a median thread-like longitudinal 
&roove both dorsally and ventrally in the plane of sym- 
metry; endocarp base and apex each with a pair of 
elliptical apertures (3-4 x 5-7 mm in diameter, located 
On either side of the plane of symmetry) opening into 
а System of lacunae; endocarp wall 5-6 mm thick, 
enclosing eight longitudinal, parallel lacunae, two ven- 
tral (condylar) and six lateral; lacunae rounded in trans- 
verse section, not connected across the plane of de- 
hiscence, their walls rugulose; the three parallel lacunae 
In each valve irregularly connected laterally; the la- 
Cunae united at their ends and extended outward. Loc- 
ule cast elongate, straight, boat shaped, length 21.0— 
39.3, avg. 32.7 mm (SD=4.75, n=22), width 11.5- 
18.4, avg. 14.7 mm (SD=1.8, n=22); thickness 6.9- 
13.7, avg. 10.0 mm (SD=1.9, п=22); acute to apiculate 
at the apex, somewhat less acute at the base, dorsal 
Surface convex, slightly and irregularly sculptured, with 
4 median longitudinal keel; angular in transverse sec- 
tion, each straight segment of the aggregately convex 
Profile opposite a lacuna in the endocarp wall; ventral 
Surface shallowly concave with a median longitudinal 
keel; placental scar median, subapical on the ventral 
Side, 

Specimens.—Syntypes of Scott (1954): UM 29920- 
29922, Hypotypes: OMSI Pb258, UCMP 10638, UF 
9249, USNM 354526, 354532, 354472, 354579, com- 
plete endocarps; UCMP 10639, 10640, UF 6844—6847, 
6850, 9254, USNM 354474-354476, 354499, 354500, 
354510, 354522, 354547, 354548, 354577-354581, 
locule casts; OMSI Pb243, 171, 253, USNM 354468, 
354470, 354473, 354482, 354515, 354516, 354525, 
354537, 424848, locule casts surrounded by lacunae 
Casts. 

Discussion. — Chandlera lacunosa is the largest of the 

larno menispermaceous fruits. It is present through- 
Out the Nut Beds deposit, represented by molds and 
Occasional casts of Ше endocarp (Pl. 27, figs. 10, 15), 
апа by chalcedony and/or sediment casts ofthe lacunae 
(РЇ 27, figs. 12—14) and locule (Pl. 27, figs. 17-20). 
Fractured specimens show the position of lacunae in 
Ше endocarp wall (Pl. 27, figs. 11, 16). I have not 
Observed Chandlera at other Clarno localities or else- 
Where in the fossil record. 

The boat-shaped locule casts with subapical placen- 
tation (Pl. 27, fig. 17) and straight longitudinal axis 
dicate affinity with the menispermaceous tribe Ti- 
Nosporeae. The endocarp is larger than that of any 

Nown living species of the tribe and the endocarp wall 


is thick relative to that of extant genera of the tribe. 
Scott (1954, 1956) named and recognized Chandlera 
as an extinct genus in the Tinosporeae. He noted that 
Chandlera is closest to the modern genus Parabaena 
in seed and condyle morphology, but noted that neither 
Parabaena nor any other living genus of Tinosporeae 
has endocarps with lacunae enclosed in their walls. 
Scott (1956) pointed out that the rows of spines in 
endocarps of extant taxa such as Parabaena are prob- 
ably homologous with the intralacunar partitions of 
Chandlera, and suggested that spiny endocarps evolved 
from unsculptured lacunate endocarps through loss of 
the outer wall. 

Little is known of the dispersal biology of Tinos- 
poreae. Perhaps the spiny endocarps of related modern 
genera are suitable for animal dispersal, the spines be- 
coming entangled with animal fur, whereas the smooth, 
lacunose endocarps of Chandlera may have been 
adapted for water dispersal. The system of lacunae 
within the endocarp wall is a striking feature of this 
genus, and could have provided buoyancy for water 
dispersal of the fruits. 


Genus ODONTOCARYOIDEA Scott 


Odontocaryoidea nodulosa Scott 
Plate 28, figures 1-10, Text-figure 13C 


Odontocaryoidea nodulosa Scott 1954, pp. 74-76, pl. 15, figs. 22- 
29, Scott 1956, рр. 77-79, fig. 2а-с. 


Description.— Fruit unilocular, single-seeded; endo- 
carp oblong, bilaterally symmetrical, pointed apically, 
truncate basally, crescent-shaped in cross section, length 
17.5-28.0, avg. 22.0 mm (SD=3.2, n=11), width 9.0- 
13.9, avg. 11.0 mm (SD- 1.36, n=11), thickness 6.0- 
10.4, avg. 8.4 mm (SD- 1.3, n=11), base with a pair 
of rounded cavities coalesced across the plane of sym- 
metry, apex with a pair of rounded cavities on either 
side of the plane of symmetry; dorsal surface convex, 
finely scabrate to smooth, ventral side hollowed out 
like a boat, with fine transverse striations; a median 
longitudinal keel running along the dorsal and ventral 
surfaces in the plane of symmetry is particularly prom- 
inent in the ventral hollow, indicating a bilobed con- 
dyle; thickness of lateral limb 3—4 mm; endocarp wall 
1.3 mm thick. Locule cast elongate, straight, boat 
shaped, length 12.9—24.1, avg. 19.8 mm (50-2.4, 
n—52), width 8.1-13.0, avg. 10.1 mm (SD- 1.1, n=53), 
thickness 5.6-8.3, avg. 7.0 mm (SD=0.67, n=46), 
pointed apically, flat basally, dorsal surface convex, 
smooth, ventral surface concave and transversely ru- 
minate with a grooved median keel over the dorsal and 
ventral surfaces, placental scar median, subapical on 
the ventral side. 

Specimens.—Syntypes: UM 29915, 29916. Hypo- 


68 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 69 


types: UF 6869, 6872, 6874, 6877, USNM 354438, 
354453, 354456, 354461, 424850, relatively complete 
endocarps; HU 60021, 60020, OMSI Pb172 (3 spec- 
Imens), UCMP 10641-10644, UF 6854-6868, 6870- 
6876, 6878-6880, UM 29917, 29918, USNM 354441- 
354455, 354457-354460, 354462-354467, isolated 
locule casts. 

Discussion.— Odontocaryoidea nodulosa is the most 
abundant moonseed in the Nut Beds and occurs 
throughout the vertical and lateral extent of the de- 
Posit. It is represented by locule casts (Pl. 28, figs. 5— 
9) and occasional casts of the complete endocarp (Pl. 
28, figs. 1-4). The locule casts are usually composed 
Of white chalcedony, occasionally of siltstone or sand- 
Stone, and sometimes are surrounded by the carbo- 
haceous or permineralized remains of the thick en- 
docarp wall without lacunae (Pl. 28, figs. 5, 10). The 
locule casts are readily distinguished from those of 
Chandlera lacunosa by the truncate base and smaller 
Size. 

This species was described by Scott (1954) as a new 
genus in the Menispermaceae, tribe Tinosporeae. He 
Noted that genera with endocarps that are straight or 
Only slightly curved from base to apex occur in the 
tribes Peniantheae, Fibraureae, and Tinosporeae, but 
Observed that “Straight endocarps possessing broad, 
deep condyles are found only in the Tinosporeae.” As 
15 name implies, this genus resembles the modern 
Odontocarya Miers, which occurs from the West Indies 
and Panama to the Amazon Basin. However, endo- 
Carps of the modern genus differ in being smaller, and 
thinner-walled. Odontocaryoidea differ from known 
living genera by the possession of the paired rounded 
Cavities at the base and apex of the endocarp. Scott 
(1956) showed how the tridentate apex and base of 
Odontocarya endocarps could be derived by reduction 
from Odontocaryoidea-like structure. 


Genus CALYCOCARPUM Nutt. ex Torr. et Gray. 


Calycocarpum crassicrustae sp. nov. 
Plate 28, figures 11-17; Plate 29, figure 1; 
Text-figure 13A 


Etymology.— Crassus (L = thick) + crusta (L = shell), 
referring to the thick wall of the endocarp. 


Description.—Fruit unilocular, single-seeded, en- 
docarp subhemispherical, cupshaped, bilaterally sym- 
metrical, elliptical to nearly circular in dorsiventral 
view, rounded apically and basally, length 19.0 mm, 
width 13.8 mm, thickness 9.0 mm; surface smooth, 
except along the ventral rim, which is rugulate; with a 
longitudinal keel in the plane of symmetry running 
along the concave ventral and less prominently on the 
convex dorsal surface of the endocarp; endocarp wall 
0.6 (adjacent to the condyle) to 1.2 mm (along dorsal 
side) thick, without lacunae, composed of fibers 9—20 
um in diameter, more than 250 um long, mostly per- 
iclinal, stratified in layers of varying orientation. Loc- 
ule cast subhemispherical, cupshaped, bilaterally sym- 
metrical, length 10.8-18.2, avg. 14.6 mm (SD=1.8, 
n=19), width 8.8—15 avg. 12.5 mm (SD=1.6, n=19), 
thickness 4.0-10.8, avg. 8.0 mm (SD=1.5, n=19), sur- 
face smooth even along the ventral rim, with a median 
longitudinal keel in the plane of symmetry developed 
both dorsally and ventrally; placental scar median, 
subapical on the ventral surface. 

Specimens.—Holotype: UF 6769, permineralized 
endocarp. Paratypes: USNM 424849, complete en- 
docarp cast; UCMP 10564, permineralized endocarp; 
OMSI Pb173 (2 specimens), 471, 1838, UCMP 10565- 
10567, 10746, UF 6770-6779, 6781-6787, USNM 
354161, 354504, 354505, 424841-424846, 354503 
(three incomplete specimens), locule casts. 

Discussion.—This species was recovered from silt- 
stone of the lower part of Face 3 as well as from the 
upper strata of the Nut Beds. It is most commonly 
preserved as chalcedony locule casts (Pl. 28, figs. 13, 
14, 17) that occur loose within the sediment mold of 
the endocarp. Occasional endocarp casts (Pl. 28, figs. 
11, 12) and permineralizations also occur. The holo- 
type is a permineralized endocarp (Pl. 28, figs. 15, 16) 
that has been sectioned to reveal internal anatomy (Pl. 
29, fig. 1). 

This fossil species conforms to the Tinosporeae in 
the large ventral condyle, subapical placenta and me- 
dian keel. In size and morphology these fruits closely 
resemble those of extant Calycocarpum. Even the ru- 
gulate sculpture of the ventral rim of the endocarp (Pl. 
28, fig. 12), and contrasting smooth ventral rim of the 
locule cast (Pl. 28, fig. 14), corresponds to the condition 


~ 


Text-figure 13.— Fruits of Menispermaceae, tribe Tinosporeae, from the Nut Beds. A, Calycocarpum crassicrustae sp. nov., endocarp ventral 
View, locule cast apical view, endocarp in transverse section. B, Chandlera lacunosa Scott, endocarp, ventral view (essentially identical to 
dorsal view), apical view showing pair of apertures, transverse section. C, Odontocaryoidea nodulosa Scott, endocarp, ventral view, apical 
View, and in transverse section. D, Tinospora elongata sp. nov., endocarp, ventral view, with portion of wall removed, revealing smooth 
Surface, and same in transverse section. E, Curvitinospora formanii sp. nov., locule cast in transverse section, ventral view, dorsal view, and 
latera] view. F, Tinospora hardmanae sp. nov., locule cast, ventral view, and in transverse section with portion of spiny endocarp reconstructed 

ased upon incomplete surface molds. С, Atriaecarpum clarnense sp. nov., locule cast ventral view, dorsal view and in transverse section. Н, 
Tinomiscoidea occidentalis sp. nov. locule cast apical view, ventral view and dorsal view. Scale bars — 1 cm. 


in the extant species. However, the fossil differs from 
extant C. /yonii (Pursh) Gray in having a thicker en- 
docarp wall. The endocarp wall of the extant genus is 
about % as thick as that of the fossil (cf. Pl. 28, figs. 
16 and 18). 

Calycocarpum crassicrustae more closely resembles 
extant C. lyonii than it does *Calycocarpum? jenkensii 
Chandler" from the London Clay (Chandler, 1961b). 
The latter is smaller and shallower than the Clarno and 
extant species, and probably represents another genus 
of the Tinosporeae. The only living species, Calyco- 
carpum lyonii, is a twining, scrambling, deciduous li- 
ana distributed in alluvial woodlands, adjacent wood- 
ed slopes and ravines in southeastern North America. 


Genus TINOSPORA Miers 


Tinospora is a genus with about 32 extant species of 
lianas distributed in tropical Africa, S.E. Asia, Indom- 
alesia, Australia and the Pacific. Referring to the habit 
of three species common in secondary vegetation in 
Thailand, in areas where forest has been destroyed or 
disturbed, Forman (1985) stated that “the exceptional 
capacity of these climbers to survive damage and to 
regenerate from detached lengths of stem encourages 
their spread in these disturbed habitats." The genus is 
represented by two species in the Nut Beds assemblage. 


Tinospora elongata sp. nov. 
Plate 29, figures 2-8; Text-figure 13D 


Etymology.— Elongatus (L — prolonged), referring 
to the relatively elongate form of the locule cast. 

Description.—Locule cast ellipsoidal, bilaterally 
symmetrical, with a medial dorsal keel in the plane of 
symmetry and a deep ventral hollow; surface smooth 
and rounded, slightly pointed apically and less so ba- 
sally, length 7.7—9.5, avg. 8.4 mm (SD=0.79, n=4), 
width 4.5-5.8, avg. 5.2 mm (SD=0.60, n=5), thickness 
3.8-4.4, avg. 4.1 mm (SD=0.31, n=3); condyle bul- 
bous, 4.8-5.2 mm long, 2.9-4.8 mm wide, intruding 
about 3.0 mm into the endocarp, with median longi- 
tudinal groove; lateral limbs of the locule cast 1.5 mm 
thick; endocarp wall thin, 0.2 mm in the condyle re- 
gion. 

Specimens.— Holotype: USNM 354502. Paratypes: 
USNM 355456, 355467, 424648, 424649, 422391, 
422392. 

Discussion.— Tinospora elongata is known from sev- 
eral locule casts. A carbonaceous film between the con- 
dyle cast and the locule (Pl. 29, figs. 3, 4) indicates that 
the endocarp wall was very thin adjacent to the con- 
dyle, however the dorsal side of the endocarp remains 
unknown. The deeply intruded bilobed condyle is best 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


seen in a transversely broken locule cast (Pl. 29, figs. 
6-8). 

Tinospora elongata closely resembles T. excavata 
Reid et Chandler and T. wilkinsonii Chandler of the 
London Clay flora (Reid and Chandler, 1933; Chandler 
1961b) in size and morphology of the condyle and 
locule cast. However, the Clarno locule casts are some- 
what more elongate in outline. It is not known if the 
endocarps of 7. elongata were tuburcled on the dorsal 
surface like those T. excavata or perhaps spined as in 
T. wilkinsonii. Among extant species, the endocarps 
range from smooth-walled to verrucate to spinose (For- 
man, 1986). 


Tinospora hardmanae sp. nov. 
Plate 29, figures 9-13; Text-figure 13F 


Menispermaceae, Bones, 1979, pl. 1, fig. 10. 


Etymology.—Named after Carol Hardman ac- 
knowledging her help with field work at the Nut Beds 
locality. 

Description.—Endocarp ellipsoidal, bilaterally sym- 
metrical, estimated dimensions: length 6.8-7.5 mm, 
width 4.2-4.8 mm, thickness 2.5-3.0 mm; concave 
ventrally with a deeply intruded smooth-walled con- 
dyle, dorsal side convex with closely spaced elongate 
spines; endocarp wall 0.6-0.8 mm thick; longitudinal 
keel in the plane of symmetry on the ventral and dorsal 
surfaces. Locule cast boat-shaped, bilaterally symmet- 
rical, elliptical in dorsiventral view, with a median 
dorsal and ventral keel in the plane of symmetry, con- 
cave ventrally, convex dorsally, rounded apically and 
basally, but with a slight apical projection from the 
keel, length 5.6-6.3, width 3.5-4.3, thickness 1.0-1.2 
mm, surface smooth; lateral limbs of locule cast 0.8 
mm thick, placental scar median, subapical on the ven- 
tral side. 

Specimens.—Holotype: UF 9255. Paratypes: UF 
9256, 9857, 9858, USNM 326715 (Bones, 1979, pl. 1, 
fig. 10), 354492, 424847. 

Discussion.—Among Clarno Tinosporeae, this spe- 
cies has the smallest fruits. Tinospora hardmanae 1s 
known from locule casts (Pl. 29, figs. 9, 10, 13) and 
fragments of endocarp molds. Although the endocarp 
itself is not preserved in these specimens, the narrow 
space between locule cast and the endocarp mold (Pl. 
29, figs. 9, 10) indicates that the endocarp was 0.6-0.8 
mm thick. Slender protrusions into the matrix at the 
basal end of the endocarp mold (Pl. 29, figs. 11, 12) 
indicate the presence of spines. The locule casts of this 
species are smaller, thinner, more elongate and тоге 
delicate than those described above as 7. elongata. 
Also, there is no evidence for spines in T. elongata. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 71 


Genus TINOMISCOIDEA Reid et Chandler 


Tinomiscoidea occidentalis sp. nov. 
Plate 30, figures 1—5; Text-figure 13H 


Etymology.— Occidens (L = of the west), denoting 
the occurrence of this species in western North Amer- 
lca. 

Description.—Locule cast shaped like a shallow boat, 
fusiform in dorsiventral view, bilaterally symmetrical, 
with a median dorsal and ventral keel in the plane of 
Symmetry, base and apex sharply pointed, length 12.0, 
16.5 mm, width 5.0, 7.1 mm, thickness 2.5, 3.4 mm; 
longitudinal axis straight, surface smooth; dorsal face 
Convex, ventral face shallowly concave, with a placen- 
tal scar subapical along the keel, 2.4 mm below the 
apex, lateral limbs of locule cast thin (ca. 1.2 mm). 

Specimens.—Holotype: USNM 422382. Paratype 
OMSI Pb926. 

Discussion.— Tinomiscoidea occidentalis is known 
only from two locule casts. The specimens closely re- 
Semble locule casts of Tinomiscoidea scaphiformis Reid 
et Chandler (1933) from the lower Eocene London Clay 
flora in size, sharply pointed base and apex and shallow 
Ventral condyle which gives a flattened appearance to 
the locule cast. The length/width proportions are also 
Similar as indicated by the ratio of 2.3 in the Nut Beds 
Specimens, compared with 1.5-2.2 in the London Clay 
Species, 

Reid and Chandler (1933) erected the fossil genus 
Tinomiscoidea to accommodate fossil endocarps and 
locule casts similar to extant Tinomiscium but differing 
by having the “ventral cavity sharply delimited by 
faceting of the endocarp wall". The Clarno locule casts 
Show rounded rather than faceted ventral surfaces (Pl. 
30, figs. 1-5), and thus differ from the generic diagnosis 
that was based on T. scaphiformis. However, taking 
Into consideration all of the other shared characters, 
It seems reasonable to place both of these species in 
the same genus. 


Genus ATRIAECARPUM Chandler 


Atriaecarpum clarnense sp. nov. 
Plate 30, figures 6-10; Text-figure 13G 


Etymology.— This species is named for the town of 

larno, Oregon. 

Description. — Locule cast obovate in face view, bi- 
laterally symmetrical, slightly curved longitudinally; 
length 9.8-12.8 mm, width 6.6-6.7 mm, thickness 2.0 
mm, apex rounded but with a short, beaked tip, base 
Tounded, obtuse, dorsal surface with 4-6 longitudinal 
Tows of about 8 to 12 closely spaced shallow, circular 
Pits; ventral side very slightly concave with a median 
longitudinal ridge, with a row of at least 10 pits on 


either side; depressions on ventral side deeper than 
those on dorsal; thickness of limb 1.7 mm. Endocarp 
wall 0.3—0.4 ит thick on ventral side. 

Specimens.— Holotype: UF 9257. Paratypes: OMSI 
Pb481, UF 9761, 9762. 

Discussion.— Although known only from a few spec- 
imens, Atriaecarpum clarnense is readily distinguished 
from other Clarno menisperms by the dimpled dorsal 
and ventral surfaces. Four of the specimens, including 
the holotype, are chalcedony locule casts. The other 
(UF 9761) is a portion of the endocarp preserved in 
sedimentary matrix, broken and exposing the inside 
of the ventral endocarp surface. In size and shallow, 
boat-like morphology it resembles Tinomiscoidea; 
however, it differs by the dimpled sculpture and the 
lack, or only faint development, of a median longitu- 
dinal keel on the dorsal surface. 

Atriaecarpum clarnense resembles the London Clay 
fossil, A. venablesii (Chandler) Chandler (1978), in 
overall shape and surface pitting of the locule cast (cf. 
Pl. 30, figs. 6-10 with Chandler 1978, pl. 4, figs. 4, 5). 
The Nut Beds specimens are larger (6.6-6.7 vs. 4.3— 
5.5 mm wide), and characters of the endocarp, such 
as the conspicuous protruberances observed in the 
London Clay species, are not known for the Clarno 
specimens. Chandler (1978) considered this extinct ge- 
nus to belong in the Fibraureae tribe rather than in the 
Tinosporeae because it lacks the deep ventral condyle 
typical of the latter. However Forman (1985) stated 
that the Fibraureae and Tinosporeae tribes may not 
be distinct because of overlap in floral and endocarp 
characters and in this treatment I have classified 
Atriaecarpum and Tinomiscoidea as representatives of 
Tinosporeae. 


Genus CURVITINOSPORA gen. nov. 


Etymology.—Curvus (L = bent) + Tinospora, refer- 
ring to the shape of the endocarp which resembles that 
of Tinospora except in being curved from base to apex. 

Type species.— Curvitinospora formanii sp. nov. 


Curvitinospora formanii sp. nov. 
Plate 31, figures 1-5; Text-figure 13E 


Menispermaceae, Bones, 1979, pl. 1, fig. 9. 


Etymology.—Named for L. L. Forman in appreci- 
ation of his contributions to the systematics of Men- 
ispermaceae. 

Description.—Locule cast bilaterally symmetrical, 
somewhat boat-shaped, but with the long axis uni- 
formly curved from base to apex resulting in a c-shaped 
lateral profile; base rounded, apex pointed; length 5.1 
mm, width 4.6 mm, thickness ca. 2.0 mm; dorsal sur- 


72 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Text-figure 14.—Fruits of Menispermaceae, tribes Cocsinieae and Menispermeae from the Nut Beds. A, Tribe Cocscinieae: Anamirta 
leiocarpa sp. nov., endocarp in longitudinal view showing dorsal keel, and longitudinal section showing hollow, bilobed condyle about which 
the locule is curved. B-F. Tribe Menispermeae. B, Diploclisia auriformis (Hollick) comb. nov., endocarp lateral view and front view. С, 
Eohypserpa scottii sp. nov., locule cast lateral view, База! view. D, Thanikaimonia geniculata gen. et sp. nov., locule cast, lateral view, front 
view. E, Palaeosinomenium venablesii Chandler, endocarp lateral view, front view. F, Davisicarpum limacioides sp. nov., endocarp lateral 


view, front view, longitudinal section. Scale bars = | cm. 


face convex, with a three-ribbed appearance formed 
by a prominent median ridge, flanked by a pair of 
longitudinal grooves, and the broad, rounded longi- 
tudinal margins; surface smooth, but with faint broad 
depressions on the dorsal and lateral faces; ventral sur- 
face concave with a median ridge raised more than the 
lateral edges, thread-like raphe at crest of the median 
ridge; placentation subapical on the ventral side; with 
a longitudinal groove on either side of the median 
ventral ridge. 

Specimen.— Holotype: USNM 326714 (Bones, 1979, 
pl. 1, fig. 9). 

Discussion.—Although known only from a single 


chalcedony locule cast, this species can be placed with 
confidence in the Menispermaceae by virtue ofits boat- 
shape and subapical ventral placenta. The median keel 
forms a ridge that is more pronounced both dorsally 
(Pl. 31, fig. 1) and ventrally (Pl. 31, figs. 2, 3) than in 
the other Clarno menisperms. 

The boat-shaped morphology of the locule in Curv- 
itinospora is similar to that of many Tinosporeae. The 
curvature of the long axis (Pl. 31, figs. 4, 5) brings 10 
mind that of Hypserpa (See Forman, 1986, fig. 12 f). 
and Eohypserpa. However, the lateral profile of Curv- 
itinospora is C-shaped, whereas that of Hypserpa, with 
both limbs meeting, is O-shaped. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 73 


Tribe MENISPERMEAE 


The Menispermeae tribe (syn. Cocculeae) is char- 
acterized by curved, often horseshoe-shaped, endo- 
carps and seeds. Endocarp sculpture provides good 
taxonomic features for the tribe (Forman, 1974, 1986). 
Two genera are present in the Nut Beds assemblage, 
the fossil genus Palaeosinomenium and the living Asian 
genus Diploclisia. Each is represented by a single spe- 
Cies, each of which appears to be identical to species 
from the Eocene London Clay flora. Fruits of this tribe 
are known from the Paleocene of Germany (Mai, 1989) 
and North Dakota (Crane et al., 1990). 


Genus DAVISICARPUM Chandler 


Davisicarpum limacioides sp. nov. 
Plate 31, figures 10-14; Text-figure 14F 


Etymology.— Limacia (an extant genus of Menis- 
Permeae) + oides (Gr = like, resembling), referring to 
the similarities between this fossil and the extant genus. 

Description. —Endocarp subglobose, unilocular bi- 
laterally symmetrical with a curved dorsal horseshoe- 
Shaped ridge forming a keel in plane of symmetry, 
laterally convex with an inflated condyle; length 4.9 
mm, estimated width in plane of symmetry 5.2 mm, 
thickness across plane of symmetry (and condyle) 3.7 
mm; dorsal keel striated perpendicular to its margin 
but smooth, not fluted; locule horseshoe shaped, sit- 
uated along the dorsal margin of the fruit, containing 
а single curved seed; endocarp wall 70-80 um thick 
Where it surrounds the locule, increased to 550—600 
4m thick on the lateral flanks of the condyle; locule 
lining uniseriate, composed of rectangular cells, 7.5— 
10 um high, 7.5-15 um long, remainder of endocarp 
formed of periclinally oriented fibers 12.5 um in di- 
ameter in that part of the endocarp forming the wall 
around the locule, and of isodiametric cells, 10-70 um 
in the lateral flanks around the condyle. 

Specimen.— Holotype: USNM 435054. 

Discussion.— Davisicarpum limacioides is known 
from a single well preserved specimen that was pho- 
tographed for external morphology (Pl. 31, figs. 10- 
12) and then sectioned transversely to show internal 
Structure (РІ. 31, figs. 13, 14). In lateral view it resem- 
bles endocarps of various genera of the Menispermeae 
tribe such as Palaeosinomenium (Pl. 29, fig. 17), how- 
Ever the keel is not fluted, and in transverse view the 
latera] margins are seen to be inflated and markedly 
Convex, rather than concave. The condyle was prob- 
ably more or less hollow originally and subsequently 
filled with sediment when the fossil was deposited. 

This species is similar to Davisicarpum gibbosum 
Chandler from the London Clay (Chandler, 1961b, 
1978), but is smaller (width 5.2 vs 7.1—7.5 mm, length 


4.9 vs 6.5—7.5 mm). Chandler (1978) noted that Dav- 
isicarpum shows the closest resemblance to extant Li- 
macia. Both genera have a curved locule and an in- 
flated condyle resulting in markedly convex lateral 
margins. The structure of the Limacia endocarp in 
longitudinal section is shown by Forman (1986, fig. 
12r) and is very similar to that of D. limacioides (Pl. 
31, figs. 13, 14; Text-fig. 14F). Justifying continued 
retention of the fossil generic name Davisicarpum, 
Chandler (1978) noted a consistent morphological dif- 
ference between the London Clay species and Limacia 
that also holds true for the Clarno specimen: “Їп Li- 
macia oblonga (Wall) Miers the locular rim is contin- 
ued further towards the attachment than in Davisicar- 
pum so as to leave a small gap only between the ends 
of the limbs, whereas in Davisicarpum the gap is much 
wider." The endocarps of L. oblonga are two to three 
times larger (15 mm high and wide) than the fossil 
specimens. The other two extant species, Limacia 
scandens Lour. and L. blumei (Boerl.) Diels, have not 
been examined in relation to the fossil material; fruit- 
ing material is not currently available among the col- 
lections of L. blumei (Forman, 1986). Nevertheless, 
Davisicarpum is clearly a close relative of this living 
genus. 

Limacia has three extant species that are woody 
climbers in tropical southeastern Asia and Malesia 
(Forman, 1986). Wolfe (1977) recognized leaves of Li- 
macia from the middle Eocene Kushtaka Formation 
of Alaska. 


Genus DIPLOCLISIA Miers 


Diploclisia auriformis (Hollick) comb. nov. 
Plate 30, figures 11—14; Text-figure 14B 
Carpolithes auriformis Hollick 1936, p. 170, pl. 120, fig. 7. 


Diploclisia bognorensis Chandler 19615, p. 161, pl. 16, fig. 14-17. 
Dipoclisia, Bones, 1979, pl. 2, figs. 2, 3. 


Description. —Fruit unilocular, single-seeded, en- 
docarp laterally compressed in plane of bisymmetry, 
obliquely obovate in lateral view with broad-rounded 
dorsal surface and acute-rounded base; length 6.3-8.0, 
avg. 7.0 mm (SD=0.75, n=5), width 4.0-6.8, avg. 4.9 
mm (SD=1.20, n=5), thickness 1.4-1.5; avg. 1.47 mm 
(SD=0.60, n=5); with a keel running along the dorsal 
surface in the plane of symmetry; both lateral margins 
bearing a horseshoe-shaped ridge separating dorsal and 
ventral ornamentation; dorsal side of the ridge marked 
with 25 to 30 radially aligned ribs alternating with 
rounded pits or grooves; inner or ventral flank of the 
ridge smooth and concave, with an obliquely oriented, 
elliptical foramen. Locule cast conforming in mor- 
phology and ornamentation to endocarp, horseshoe- 
shaped with approximately equal limbs. 


74 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Specimens.—UCMP 10558, 10559, UF 6484, 6485, 
6486, USNM 312648 (Bones, 1979, pl. 2, fig. 3), 312749 
(Bones, 1979, pl. 2, fig. 2), 354508, 354571, 354573, 
446069. 

Discussion. — This species is represented in the Nut 
Beds by carbonaceous endocarp compressions (Pl. 30, 
figs. 12, 13) and chalcedony locule casts (Pl. 30, figs. 
11, 14). The compressions closely resemble those of 
Carpolithes auriformis Hollick from a Paleogene lo- 
cality on the south shore of King Salmon Lake, Alaska 
Peninsula (Hollick, 1936, pl. 120, fig. 7), and the casts 
closely resemble those described as Diploclisia bog- 
norensis from the London Clay (Chandler, 1961b). 
These occurrences appear to represent a single mor- 
phological species. Endocarps with the horseshoe 
shaped locule and fluted dorsal surface are diagnostic 
of the Menispermaceae, tribe Menispermeae. 

Diploclisia occurs today in East Asia, China and Bur- 
ma to India and the Malay Peninsula. In reference to 
specimens of this species from the London Clay flora 
Chandler (1961b, р. 162) stated that “The form and 
structure are so closely comparable with the living Di- 
ploclisia that there can be по doubt of the relationship.” 
Among the most similar extant species are D. affinis 
Diels and D. chinensis Merrill. Endocarps of D. affinis 
(specimen examined E.H. Wilson 2286 from western 
Hupeh, China) are particularly similar in size, shape, 
number of ribs, and the smooth ventral surface to the 
fossil species. 


Genus EOHYPSERPA Reid et Chandler 


Eohypserpa scottii sp. nov. 
Plate 29, figures 14-16; Text-figure 14C 


Etymology.—This species is named for Richard A. 
Scott, recognizing the importance of his contributions 
to fossil Menispermaceae. 

Description.—Fruit subglobose, nearly circular in 
lateral view; locule cast bilaterally symmetrical, smooth, 
prominently curved with base and apex nearly touch- 
ing, width across plane of symmetry 7.3 mm, diameter 
in plane of symmetry 8.5-11.5 mm; median keel in 
the plane of symmetry on the dorsal (and ventral?) 
surface becoming more prominent at the apical end, 
apical limb pointed, basal limb blunt-rounded; with a 
central circular aperture 2-2.5 mm in diameter on each 
lateral face, about which the locule is curved. 

Specimens.— Holotype: USNM 354506. Paratype: 
USNM 446075. 

Discussion.— Eohypserpa is represented by two loc- 
ule casts from the Nut Beds. Although the endocarp 
itself is not preserved, the locule morphology is very 
distinctive and is similar that of extant Hypserpa (see 
Forman, 1986, fig. 12). In both there is a central pair 


of lateral apertures in the endocarp about which the 
locule is curved. Locule casts of Eohypserpa зсоий 
resemble those of the London clay species, Е. parsonii 
Reid et Chandler (1933), in general form, smooth sur- 
face and size. The London Clay species was diagnosed 
largely on the basis of endocarp features that are not 
preserved in the Nut Beds specimens. 

Reid and Chandler (1933) observed the striking sim- 
ilarity between the London Clay endocarps and those 
of extant Hypserpa, but reported several differences 
that led them to place the fossils in an extinct genus. 
These differences include the following: 1) Reid and 
Chandler stated that the fruits of extant Hypserpa are 
more compressed laterally, whereas those of the Lon- 
don Clay are typically more globose [however, there 
is overlap in this feature]; 2) In Hypserpa the curvature 
of the carpel is less symmetrical than in the London 
Clay fossils; 3) In Eohypserpa the cross section of the 
locule is more elongate but more sharply curved at the 
cusps than in the living genus, and the lateral cavities 
are relatively larger; 4) The fossil endocarps from the 
London Clay are transversely ribbed, but not tubercled 
or angular as in Hypserpa. 

It is not known whether the Clarno species had a 
transversely striate endocarp as in the London Clay 
genus, or a verrucate endocarp as in extant species of 
Hypserpa. However, the available characters including 
symmetry of the locule, curvature and size of the lateral 
cavities, and smoothness of the locule support place- 
ment of the Clarno species in Eohypserpa rather than 
Hypserpa. 


Genus PALAEOSINOMENIUM Chandler 


Palaeosinomenium venablesii Chandler 
Plate 30, figures 15-20; Text-figure 14E 


Palaeosinomenium venablesii Chandler 1961b, pp. 159-160, pl. 16, 
figs. 9-13. 


Description.—Fruit unilocular, single-seeded, en- 
docarp laterally compressed in plane of bisymmetry, 
obliquely ovate in lateral view, with broad-rounded 
dorsal margin and straight ventral margin, endocarp 
length 2.1, 3.5, 3.7 mm, width 2.3, 4.4, 4.3 mm, thick- 
ness 1.0, 1.9, 1.9 mm; dorsal margin of endocarp with 
a prominent median keel in the plane of symmetry 
(0.4-0.5 mm high); both lateral faces with a broad, 
shallow concavity rimmed by a C-shaped ridge sepa- 
rating dorsal and ventral ornamentation; each lateral 
face with 18-21 radially aligned ridges alternating with 
rounded grooves that are developed both ventrally and 
dorsally; an obliquely oriented narrow foramen is sit- 
uated medially near the base. Locule cast obliquely 
C-shaped with the two arms of unequal length, one of 
the arms blunt-tipped, the other tapered to a point, 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 75 


Not as sharply sculptured as the endocarp, fluted dor- 
sally with 14-17 depressions, smooth ventrally; length 
2.0-4.1, avg. 2.9 mm (SD=0.61, n=41), width 2.3-4.3, 
avg. 3.3 mm (SD=0.56, n=41), thickness 0.81-2.60, 
avg. 1.33 mm (SD=0.37, n=41), ventral margin mark- 
edly concave, dorsal margin convex. 

Specimens.—USNM 354461, 424815, 424826, en- 
docarp casts; HU 60018, 60019, UCMP 10560-10563, 
UF 6487-6491, 6737, USNM 354494 (17 specimens), 
354489 (6 specimens), 354491 (5 specimens), 354493, 
354514, 354559-355567, 424812-424814, 424816- 
424840, 435101, locule casts. 

Discussion. — This species is known from the Nut 
Beds mostly from isolated chalcedony locule casts (Pl. 
30, figs. 19, 20). Molds in the matrix from which the 
locule casts were removed show the impression of the 
endocarp surface (Pl. 30, fig. 15). Rarely, specimens 
are preserved as endocarp casts showing the exterior 
Sculpture (Pl. 30, figs. 16-18). In all observable char- 
acters, the Clarno specimens are indistinguishable from 
the London Clay specimens named Palaeosinomenium 
venablesii Chandler, although details of internal anat- 
Оту described by Chandler (1961b) are not preserved 
In the Clarno specimens. 

Palaeosinomenium endocarps and locule casts bear 
Some resemblance to those of Diploclisia described 
above, reflecting their common affinities within the 
tribe Menispermeae. However, P. venablesii is readily 
distinguished from D. auriformis by a smaller length/ 
Width ratio, by the unequal length of the limbs, and 

Y the presence of ventral sculpture. Similar extant 
8епега in the Menispermeae include Menispermum and 
Sinomenium. But according to Chandler (19611), they 
аге distinguished from the fossil by being less oblique 
ànd having shorter, rounder foramina. Cocculus en- 
docarps are more rounded and inflated than those of 
the fossil, and Cissampelos has a median groove in- 
Stead of a median keel. 


Genus THANIKAIMONIA gen. nov. 


Etymology.— This genus is named in memory of G. 

hanikaimoni, recognizing his contributions to the 
Morphology and systematics of Menispermaceae. 

Type species. — Thanikaimonia geniculata sp. nov. 


Thanikaimonia geniculata sp. nov. 
Plate 31, figures 6-9; Text-figure 14D 


Etymology.— Geniculate (L — like the bent knee), 
Teferring to the strongly curved locule cast. 

Description .— Locule cast obovate, horseshoe- 
Shaped, bilaterally symmetrical, length 5.3 mm, width 
m plane of symmetry 4.3 mm, thickness across plane 
ЈЕ Symmetry 4.0 mm, dorsal and ventral surfaces 
Smooth, with a median dorsal keel in the plane of 


symmetry; both lateral faces with a prominent median 
concavity about which the seed cavity is curved; limbs 
ofthe seed cavity nearly equal in length, but with one 
broader and more rounded at its extremity than the 
other, the broader one with a circular scar at its tip. 

Specimen.— Holotype: USNM 355480. 

Discussion. — This species is represented by a single 
locule cast showing a horseshoe-shaped seed cavity (Pl. 
31, figs. 7, 8) with a median dorsal keel in the plane 
of bisymmetry (Pl. 31, figs. 6, 9). In general form the 
locule cast of Thanikaimonia resembles that of Diplo- 
clisia, but it differs by the lack of surface sculpture and 
by its greater thickness across the plane of symmetry. 
Smooth locules occur among extant Menispermeae, as 
in Pachygone, but no close morphological match to 
this fossil has been found among extant genera. 


Family MUSACEAE Juss. 


The Musaceae (banana family) have three modern 
genera of old world tropical distribution: Musa L. (35 
spp.), which is restricted to Asia, Ensete Horan. (7 spp.) 
of Asia and Africa, and Musella C.Y. Wu ex H.W. Li 
(1 sp.), in southern China. Ensete was treated as a 
subgenus of Musa prior to its recognition as a genus 
by Cheesman (1947), and Musella was only recently 
segregated from Ensete (Wu, 1981). Features distin- 
guishing Ensete from Musa include persistent bracts 
and flowers, warty rather than granular pollen, and 
larger seeds with a deeply recessed hilar cavity. Musella 
differs from the other two genera by its short stems 
and congested rhizomatous growth. 


Genus ENSETE L. 


Ensete oregonense Manchester et Kress 
Plate 32, figures 1-14, 16-19 
Ensete oregonense Manchester et Kress 1993, pp. 1264-1272, figs. 
2-11, 37. 
Olacaceae, Bones, 1979, pl. 5, fig. 1. 


Description.—Seed anatropous, subfusiform-ellip- 
soidal when complete, circular to subangular in trans- 
verse section, base rounded, apex truncate; length of 
complete seed 10.0-12.0, avg. 10.8 mm (SD=0.96, 
n=4), width 5.3-7.2, avg. 5.8 mm (SD=0.43, n=15), 
with a wide hilar depression, and an operculum (mi- 
cropylar plug) about 1.2 mm in diameter; hilar rim ca. 
1 mm high; outer surface of seed smooth to finely 
striate longitudinally; seed coat of two layers, each 200— 
250 um thick and composed of longitudinal fibers, the 
inner layer diverging from the outer seed coat near the 
base and apex of the seed, delimiting a chalazal mass 
at one end, a micropylar collar at the other, and a 
central barrel-shaped embryo/endosperm chamber; 
embryo small, 2.5 mm long, straight and bulbous, as- 
cending from the channel delimited by the micropylar 


76 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


collar into the central chamber. Length of specimens 
with hilar rim broken away 6.1-10.0, avg. 7.5 mm 
(SD=0.98, n=15); Length of central chamber (from 
casts missing both the chalazal mass and hilar rim) 
4.3-8.5, avg. 6.0 mm (SD=0.84, n=47), width 5.3-7.2, 
avg. 5.8 mm (SD=0.43, n=15). 

Specimens.—Holotype: UF 6621. Paratypes: HU 
59915, 60039, 60041, OMSI Pb1061, 1314, 1842, 
UCMP 10688-10695, UF 6611-6620, 6622-6642, 
6726, 9739, USNM 326732 (Bones, 1979, pl. 5, fig. 
1), 354608-354613, 354615-354617, 354621-354625, 
354626, 424620, 434966-434982, UWBM 35274, 
36056, 36080. 

Discussion. — Seeds of Ensete are especially common 
in the Nut Beds. They occur in the basal leaf layer in 
Face 4, and are common at the top of Face 3. The 
seeds are preserved in different fashions. Most are chal- 
cedony infillings of the central chamber of the seed. 
These are barrel-shaped casts with a depression and 
circular mark at both ends, sometimes with adhering 
remnants of tegmen and testa (Pl. 32, figs. 3-5, 8, 9). 
Occasional specimens are more complete, with the 
chalazal region intact showing the rounded base. In 
these the profile of the seed is faintly constricted in the 
position corresponding to the septum separating the 
central chamber and chalazal mass (Pl. 32 figs. 6, 10). 
The seeds usually are preserved with the operculum 
detached, exposing the micropylar collar. Longitudinal 
sections reveal that the seeds are bitegmic (Pl. 32, fig. 
14), with the tegmen diverging from the testa near the 
base and apex ofthe seed, delimiting the chalazal mass 
at one end and the micropylar collar at the other, and 
a central barrel-shaped endosperm/embryo chamber. 
A small straight embryo protrudes into the endosperm 
chamber from the micropylar end (Pl. 32, figs. 12, 13). 
A few specimens show a conical projection at the mi- 
cropylar end (Pl. 32, fig. 4) corresponding in position 
to the *nucellar pad" of McGahan (1961). In a few 
cases the operculum is preserved (Pl. 32, fig. 16). 

The affinities of these and other fossil and extant 
seeds of Zingiberales were treated in detail by Man- 
chester and Kress (1993). Ensete oregonense seeds con- 
form to those of the Musaceae 1n the thick seed coat, 
well-defined micropylar collar, operculum, and a sep- 
tum dividing the embryo chamber from the chalazal 
mass. Ensete is differentiated from Musa by its ten- 
dency for larger seeds (1 to 2.5 cm), and an enlarged 
hilar depression. The fossil seeds are at the high end 
of the size range for extant Musa seeds and at the low 
end ofthe size range for extant Ensete seeds. The fossils 
clearly belong to Ensete because of the enlarged hilar 
depression (cf. pl. 32, fig. 15). 

This species is the only known record of Musaceae 
based upon seeds from North America. Although no 


musaceous leaves have been confirmed from the Nut 
Beds, it is likely that there are significant taphonomic 
biases against transport of large, persistent leaves to 
sites of deposition, whereas the seeds could easily have 
been carried by birds and/or water runoff. The confir- 
mation of Ensete seeds in the North American Tertiary 
lends credence to the identification of the family based 
on leaves of Musophyllum complicatum Lesquereux 
from the Eocene Green River Formation of Wyoming 
(Lesquereux, 1878, pl. 15, fig. 1—6). 


Family PALMAE Juss. 


The Palmae (palm family) are represented in the Nut 
Beds by silicified stems and petioles and by Sabal-like 
leaf impressions (Manchester, 1981) as well as by seeds. 
Two species of Sabal are recognizable on the basis of 
seeds. Previous reports of palm seeds from the Nut 
Beds (e.g., Bones, 1979) were based upon specimens 
that I have dismissed from the Palmae and treated 
here as Coryloides in the Betulaceae. 


Genus SABAL Adans. 


Sabal is a genus of 15 extant species distributed in 
the southeastern United States and Mexico, the West 
Indies and northern South America (Zona, 1990). The 
genus has also been confirmed on the basis of cuticu- 
larly preserved costapalmate leaves from the Eocene 
of Tennessee (Daghlian, 1978), and from seed remains 
from the Tertiary of Europe (Mai, 1976). The two spe- 
cies of Sabal seeds from the Nut Beds are considered 
below. 


Sabal bracknellense (Chandler) Mai 
Plate 33, figures 1-3 
Palmospermum bracknellense Chandler 1961b, pp. 125-126, pl. 13, 
figs. 15, 16. 
Sabal bracknellense (Chandler) Mai 1976, pp. 104—105, pl. 2, figs 
5-9, 
Palmae, Bones, 1979, pl. 1, fig. 3. 


Description.—Seed cast globose to oblate, circular in 
dorsiventral plane section, rounded dorsally, truncate 
ventrally, height and width 3.8—5.3, avg. 4.3 mm 
(SD=0.48, п- 14), dorsiventral thickness 4.0-6.8, avg. 
4.7 mm (SD=0.66, n=15); ventral truncation circular; 
shallowly depressed, with a central circular raised area 
1.2-2.0 mm in diameter; sometimes with a shallow 
longitudinal groove passing over the rim of the ventral 
truncation; rim around the truncation rounded, faintly 
ruminate; otherwise, seed surface smooth, occasionally 
with a small circular dome-like area on the surface 
representing protrusion of the embryo. 

Specimens.—UF 8484-8491, USNM 354662 (7 
specimens), UCMP 10720-10727. 

Discussion.— This is the more common species of 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 77 


Sabal in the Nut Beds assemblage. Although preserved 
Only as internal molds of the seed coat, without details 
of anatomy, these specimens correspond very closely 
in size and morphology to seeds of Sabal bracknellense 
(Chandler) Mai from the Eocene London Clay (Chan- 
dler, 1961b) and Geiseltal (Mai, 1976) floras. The cir- 
Cular dorsiventral outline, and ventral depression with 
Central hilum (Pl. 33, figs. 1-3) are characters diag- 
Nostic of extant Sabal. Referring to Palmospermum 
Jenkinsii, P. bracknellense and P. cooperi from the Lon- 
don Clay, Mai (1976, p. 105) questioned why Chandler 
included these species in a form-genus for palm seeds 
Of unknown affinities: “The characteristics of all these 
Species speak very distinctly for their placement in the 
genus Sabal Adans.” 


Sabal jenkinsii (Reid et Chandler) 
comb. nov. 
Plate 33, figures 4—8 


Palmospermum jenkinsi Reid et Chandler 1933, p. 110, pl. 1, figs. 
25724 


Description.—Seed cast globose to oblate, circular in 
dorsiventral view, rounded dorsally, truncate ventral- 
ly; height and width 7.3-9.0 mm, dorsiventral thick- 
Ness 5.7-7.4 mm; ventral truncation circular, shallowly 
depressed, with a central raised area 2.2 mm in di- 
ameter, from which cells radiate; with a shallow lon- 
8itudinal raphe groove passing over the rim of the 
ventral truncation; seed surface smooth, with a small 
Circular dome representing the protruding embryo vis- 
10е on some specimens; seed coat 130 um thick, of 
two layers, inner layer darker, 20-30 um thick com- 
Posed of uni- or biseriate row of rectangular cells, outer 
layer lighter, not well preserved appearing almost ge- 
latinous but with sharp boundaries, 100 um thick; with 
an additional layer of sclerenchymatous tissue some- 
ümes preserved in the ventral depression. 

Specimens. —OMSI Pb1812, UF 975 1-9754, USNM 
326713 (Bones, 1979, pl. 1, fig. 3). 

Discussion.— Sabal jenkinsii is represented in the Nut 
*ds only by a few specimens from the top of Face 3. 
П one of the specimens a thickened patch of the seed 

Coat is preserved in the ventral hollow (Pl. 33, fig.7). 

Dé specimen still partially embedded in the sedi- 
Mentary matrix was sectioned to show the bilayered 
Wall structure (Pl. 33, fig. 8). 

This species corresponds to Sabal in overall shape, 
Seed coat thickness, and configuration of the ventral 

*pression. Just as in extant seeds of the genus, the 
Seed coat is thickened in the hilar region much more 
than it is over the body ofthe seed. The main difference 
between this species and S. bracknellensis is the larger 
Size. This species conforms to the diagnosis of Pal- 


mospermum jenkinsii Reid et Chandler (1933), and 
following the suggestion of Mai (1975), is hereby trans- 
ferred to Sabal. 


Family PLATANACEAE Dumort. 


The Platanaceae are a family of one extant genus 
and about 7 species of trees. The family has an excellent 
fossil record that includes inflorescences and infruc- 
tescences extending into the mid-Cretaceous (Friis, et 
al., 1988). The Platanaceae are represented in the Nut 
Beds by wood of Plataninium (Scott and Wheeler, 
1982), leaves of Macginitiea (Manchester, 1986) and 
the three genera of platanaceous infructescences treat- 
ed below: Macginicarpa, Platanus, and the new genus 
Tanyoplatanus. 


Macginicarpa glabra Manchester 
Plate 33, figures 9-12 


Macginicarpa glabra Manchester 1986, р. 209, figs. 20-37, 44. 


Description.—Infructescence spheroidal, 9-15 mm 
in diameter, consisting of a central receptacular core 
3—6, avg. 4.5 mm in diameter surrounded by about 60 
closely packed florets; florets hypogynous, each con- 
taining five, very rarely six, apocarpous achenes arising 
from a pentagonal (rarely hexagonal) receptacle, sur- 
rounded by a perianth of 15-30 tepals; tepals extending 
as high as the achenes, 3.5 mm long, 0.6-1.0 mm wide, 
and 0.13-0.5 mm thick, outer tepals thicker than inner, 
but perianth not differentiated into petals and sepals; 
achenes free, arranged in a single whorl with ventral 
sutures contiguous about the floral midline, achene 
body ovoid to ellipsoidal, less commonly obovoid 3- 
4 mm long, 0.5-1.2 mm wide, 0.7-1.5 mm deep in 
predispersal state, 5-8 mm long and 2.5-3.0 mm wide 
or deep at maturity, with a persistent style up to 2.3 
mm long, slightly curved, with a ventral slit running 
its full length; achenes glabrous, pericarp consisting of: 
the exocarp, with a thin uniseriate layer of cutinized 
epidermal cells, the mesocarp, composed mainly of 
isodiametric parenchyma cells 35—50 um in diameter, 
and the endocarp, made up of a uniseriate layer of 
anticlinally elongate, thick-walled cells, 7-10 um wide, 
60 ит high. 

Specimens.— Holotype: UF 5153. Additional spec- 
imens: UF 5153-5159, 5161-5164, USNM 354077- 
354083, 354090-354092, 354095, 354097, 354104- 
354106, 354115, 354116, 354119-354140, 354142- 
354148, 354150, 354151, 354587. 

Discussion.— Macginicarpa glabra infructescences 
are one of the most abundant reproductive structures 
in the Nut Beds deposit. They are preserved both as 
compressions and as permineralizations. Excellent an- 
atomical details are provided by permineralized fruits 


that have been fractured (Pl. 33, fig. 9) and/or sectioned 
(Pl. 33, figs. 10-12). The same species occurs at several 
other localities in the Clarno Formation, consistently 
associated with digitately palmately lobed leaves of 
Macginitiea angustiloba. Based upon shared platana- 
ceous affinity, and co-occurrence at several localities, 
I have suggested that the M. angustiloba leaves be- 
longed to the same plant that produced Macginicarpa 
infructescences—a plant informally termed the Clarno 
plane tree (Manchester, 1986). Silicified staminate in- 
florescences (Platananthus synandrus Manchester, Pl. 
33, fig. 13), isolated stamen groups (Macginistemon 
mikanoides [MacGinitie] Manchester) and platana- 
ceous wood (Scott and Wheeler, 1982) from the Nut 
Beds are also hypothesized to correspond to the Clarno 
plane tree (Manchester, 1986). 

Macginicarpa glabra conforms to Ше Platanaceae in 
the possession of unisexual globose heads of numerous 
florets each bearing several achenes with persistent 
styles and long ventral sutures. However, the species 
differs from living species of Platanus because: 1) it 
has a relatively well developed perianth, 2) the number 
of achenes per floret is relatively constant at five, rather 
than variable within the head, 3) the achenes usually 
are ovoid rather than obovate, 4) dispersal hairs are 
lacking. The lack of dispersal hairs in achenes of Mac- 
ginicarpa suggests that these fruits were not as well 
adapted for wind dispersal as those of extant Platanus 
(Manchester, 1976). Infructescences of Macginicarpa 
are also known from the Early Eocene of Chalk Bluffs, 
California, Middle Eocene of Republic, Washington 
(Manchester, 1986), and from the Paleocene of Joffre 
Bridge, Alberta (Pigg and Stockey, 1991), but are not 
known from the European or Asian Tertiary. 


Genus PLATANUS L. 


Platanus hirticarpa sp. nov. 
Plate 33, figure 14; Plate 34, figures 1-4 


Etymology.— Hirtus (1. = hairy, shaggy) + karpos 
(Gr = fruit), referring to the presence of dispersal hairs 
on the fruitlets. 

Description.—Infructescence spheroidal, 8.0 mm in 
diameter, consisting of a central receptacular core ca. 
2 mm in diameter surrounded by numerous closely 
packed florets; florets hypogynous, each containing four 
(or more?), apocarpous achenes, surrounded by a peri- 
anth of about 15 tepals; tepals extending as high as the 
achenes, 2.2 mm long, 0.5-1.0 mm wide, and 20-30 
um thick, outer tepals thicker than inner, but perianth 
not differentiated into petals and sepals; achene body 
obovate to obtriangular in longitudinal outline, with a 
persistent style up to 2.3 mm long, slightly curved, 
with a ventral slit running its full length; achenes en- 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


veloped with numerous hairs that arise from the lower 
portion of the fruit, these hairs large, 24—29 шт in 
diameter, 1250-1500 um long, uniseriate-multicellu- 
lar, composed of cells 80-100 um long. 

Specimens.— Holotype: UF 5160. 

Discussion.—In contrast to Macginicarpa which is 
abundant in the Nut Beds, extant Platanus is known 
only from a single infructescence. At the time Mac- 
ginicarpa was described (Manchester, 1986), I didn’t 
know that Platanus was present as well. In addition to 
the infructescence described above, Platanus is now 
known from rare isolated achenes with the character- 
istic dispersal hairs as compression fossils from three 
other Clarno localities (West Branch Creek, White 
Cliffs, Sheep Rock Creek). 

On the basis of infructescence morphology (Pl. 33, 
fig. 14), achene shape (Pl. 34, figs. 2, 3) and presence 
of dispersal hairs (Pl. 34, fig. 4), I consider that this 
fossil belongs to the extant genus. Macginicarpa and 
the Cretaceous platanaceous infructescences known as 
Platanocarpus Friis, Crane et Pedersen (1988) differ 
from Platanus because the achenes lack dispersal hairs. 
Macginicarpa is also distinguished by ovate, rather 
than obtriangular, achenes. P. hirticarpa represents the 
earliest record of extant Platanus to be confirmed by 
fruit structure, and shows that both extant and extinct 
genera of Platanaceae coexisted in the early Tertiary. 
Platanaceous leaves that appear to represent Platanus, 
and not Macginitiea, are now known from the White 
Cliffs Clarno locality, although they have not been re- 
covered from the Nut Beds. 


Genus TANYOPLATANUS gen. nov. 


Etymology.— Tanyo (Gr = elongate) + Platanus (Gt 
= plane tree), referring to the elongate infructescences 
of this platanaceous plant. 

Type species. — Tanyoplatanus cranei sp. nov. 


Tanyoplatanus cranei sp. nov. 
Plate 34, figures 5-15 


Etymology.— The epithet is named after Peter К. 
Crane, recognizing his contributions to paleobotany 
including the fossil record of Platanaceae. 

Description. — Infructescence elongate-cylindrical, uP 
to atleast 30 mm long, 5.0, 10.0, 11.0 mm in diameter; 
consisting of a straight central axis 1.6-2.8 mm in di- 
ameter surrounded by numerous closely packed florets; 
florets hypogynous, each containing three to four ap“ 
ocarpous achenes, surrounded by a perianth of about 
five to ten thin tepals; tepals extending as high as the 
achenes, perianth not differentiated into petals and 5е- 
pals; achene body ovate to less commonly obovate in 
longitudinal outline, up to 1.1 mm in diameter, 3.4 
mm long, tapered to a persistent style, with a ventral 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 79 


slit running its full length; numerous hairs arising from 
the lower portion of the fruit and receptacle; hairs large, 
20 um in diameter, more than 500 um long, uniseriate- 
Multicellular, the cells 80-100 um long. Achene anat- 
Omy: most of the achenes infertile, lacking locule de- 
velopment, composed of angular parenchyma cells; oc- 
Casionally achenes fertile, with a single locule and seed; 
locule lining comprising a uniseriate layer of tall co- 
lumnar cells, 50-80 um high, 10-12 um wide, sur- 
rounded by mesocarp of angular parenchyma cells 40- 
70 um in diameter, with about three vascular bundles, 
схосагр uniseriate layer of small rectangular cells about 
5 um thick and 10-12 um long. 

Specimens.—Holotype: USNM 424876. Paratypes: 
OMSI Pb923, UF 5671. 

Discussion.— Tanyoplatanus is represented by three 
Permineralized specimens from the Nut Beds, and by 
a few compression specimens from the West Branch 
Creek and White Cliffs Clarno localities. The general 
арреагепсе is that of an elongate bottle brush or test 
tube brush with numerous florets arranged about an 
Clongate axis. Although general aspects of the mor- 
Phology are apparent from longitudinally fractured 
Specimens (Pl. 34, figs. 5, 7), it was necessary to prepare 
Serial transverse (Pl. 34, fig. 6) and longitudinal sec- 
tions (PI. 34, figs. 7-11, 13-15) and acetate peels of 
Ше infructescences to document anatomical details. 

The infructescences of all previously described fossil 
and extant Platanaceae are more or less globose. Tan- 
Yoplatanus compares favorably with Platanus in the 
Morphology and anatomy of achenes and in the ar- 
rangement of florets on an expanded receptacular core. 
t differs by the elongated nature of the infructescence 
and by the relatively small number of achenes per floret 
(3-4 Vs. 4—9). Although this is the first formal descrip- 
Чоп of the taxon, Ше genus has been illustrated pre- 
Viously and described as “elongate infructescence” 
Crane et al., 1990) from the Paleocene of North Da- 

Оїа, and I have collected it at several localities of the 
- aleocene Fort Union Formation in Wyoming. Both 
Ш these Paleocene localities and at the Clarno West 
Branch and White Cliffs localities, compound leaves 
ofA verrhoites affinis (Newberry) Hickey co-occur, and 
lt is possible that these Sapindopsis-like leaves and 
Infructescences were borne by the same taxon. 

Unlike Macginicarpa, and like Platanus, the fruits 
= Tanyoplatanus bear numerous dispersal hairs (Pl. 
34, fig. 14). The perianth is not as well developed as 
mM acginicarpa, and the number of fruits per floret is 
Only three to four (Pl. 34, figs. 10, 11, 13), rather than 

ve. Anatomically, the achenes of Tanyoplatanus and 
асвіпісағра are very similar, with an endocarp of 
columnar sclereids, a mesocarp of angular parenchyma 
> and a thin uniseriate cutinized exocarp (Pl. 34, 
8. 15). 


A single elongate staminate catkin (UF 9342) was 
recovered from the Nut Beds deposit that may also 
correspond to Tanyoplatanus. It has two stamens per 
floret with short filaments, capitate connectives and 
small tricolpate reticulate pollen. 


Family ROSACEAE Juss. 


The Rosaceae, or rose family, includes about 107 
genera and 3000 species of trees shrubs and herbs. The 
family does not appear to be very important in the 
Nut Beds flora, being represented only by the genus 
Prunus. Two species of Prunus are recognized below 
on the basis of fruit types, and the same genus is also 
represented in the Nut Beds by silicified wood (E. 
Wheeler, pers. comm. 1991). 


Genus PRUNUS L. 


Prunus olsonii sp. nov. 
Plate 35, figures 1-10 


Etymology.—This species is named after Duane Ol- 
son, recognizing his help with field work in the Clarno 
Formation. 

Description.—Fruit unilocular, single-seeded; en- 
docarp pyriform, bilaterally symmetrical, rounded or 
rounded-truncate basally, tapering apically to an acute 
apex, with an apical keel in the plane of symmetry, 
length 6.6-6.8 mm, width 5.3-5.9 mm, thickness 4.5- 
5.3 mm, endocarp wall 0.35-0.65 mm thick, composed 
of, in succession: 1) a layer of fibers (five to eight cells 
and about 60 um thick); inner part of the fibrous layer 
with fibers mostly oriented horizontally, the rest of the 
layer with fibers mostly oriented longitudinally, 2) the 
outer layer (0.2 mm) thick, composed of polygonal 
sclereids 125-550 um in diameter with interspersed 
idioblasts about 20 um in diameter, each with a single 
rhomboidal crystal. Locule cast similar in shape to the 
endocarp, smooth, usually with longitudinal raphe 
groove aligned with the apical keel, running from the 
apex to the chalazal truncation, length 5.8-6.0 mm, 
width 4.5-5.0 mm, thickness across plane of symmetry 
4.1-4.3 mm. Seed coat forming a uniseriate layer of 
thin-walled rectangular cells 30 um high and 50-90 um 
wide. 

Specimens.—Holotype: UF 9262, permineralized 
endocarp. Paratypes: UF 9748-9750, permineralized 
endocarps; UF 6460, 6461, USNM 354999, locule 
casts. 

Discussion.— Prunus olsonii is represented by a few 
permineralized specimens and several locule casts. The 
locule casts show distinctive features such as pyriform 
shape (Pl. 35, figs. 1, 4), circular chalaza (Pl. 35, figs. 
2, 3, 6, 7) and apical keel (Pl. 35, fig. 5). The main 
thickness of the endocarp is composed of sclereids with 


80 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Text-figure 15. — Transverse section of Emmenopterys dilcheri sp. 
nov., showing two carpellate fruit with seeds on axile placentae. 


interspersed crystal-bearing idioblasts (Pl. 35, fig. 10). 
Prunoideae endocarps have a good fossil record in the 
European Tertiary (Mai, 1984), and have also been 
described from the Eocene Princeton chert of British 
Columbia (Cevallos-Ferriz and Stockey, 1991). Cev- 
allos-Ferriz and Stockey (1991) recognized three types 
of Prunus endocarps from the Princeton chert which 
they named informally types 1 to 3. Similar idioblasts 
and crystals were described in the endocarp that they 
designated Prunus type 1 (Cevallos-Ferriz and Stockey, 
1991). 


Prunus weinsteinii sp. nov. 
Plate 35, figures 11-15 


Etymology.—This species is named after Michael 
Weinstein, recognizing his invaluable assistance with 
field work in the Clarno Formation. 

Description.—Fruit unilocular, single-seeded, en- 
docarp cherry pit-shaped, bilaterally symmetrical, el- 
liptical in cross section, ovate in longitudinal view, 
with a longitudinal keel running along the margin in 
the plane of symmetry; base rounded, apex pointed; 
length 10.4 mm, width 8.6 mm, thickness 6.2 mm, 
surface gently rugulate; endocarp wall 0.8-1.2 mm thick, 
thickest in the region of the longitudinal keel, com- 
posed of polygonal, flat-sided, angular sclereids 25-38 
um in diameter (these cells becoming thinner walled 
and somewhat larger toward periphery) without ob- 
vious vascular bundles and with scattered idioblasts 
20-30 um in diameter each with a rhomboidal crystal; 
with an abscission/suture plane running from base to 
apex in the plane of symmetry along the ventral mar- 
gin; a pair of vascular bundles straddling the abscission 
plane and running along the exterior of the endocarp; 
locule rounded, smooth lining. Seed coat with a thin 
uni-to biseriate outer layer 30 um thick made up of 
horizontally elongate cells; preserved in dark color. 

Specimen.— Holotype: UF 6800. 


Discussion.— Although Prunus weinsteinii is known 
only from a single specimen, the anatomical preser- 
vation is excellent (Pl. 35, figs. 13-15). This species 15 
distinguished from other unilocular fruits of similar 
size from the Nut Beds by the pronounced keel in the 
plane of symmetry and uniformity and compactness 
of the cells making up Ше endocarp. The restriction of 
the dehiscence plane to the ventral side (and absence 
on the dorsal side) is consistent with Prunoideae and 
distinguishes it from morphologically similar endo- 
carps of Icacinaceae. This species differs from P. ol- 
sonii, treated above, by the larger size, thicker endo- 
carp wall, and lack of the layer of fibers lining the 
locule. In overall shape, wall thickness and anatomy 
the endocarp closely matches that of domesticated 
cherry, Prunus avium (L.)L. 


Family RUBIACEAE Juss. 


The Rubiaceae are a large family of about 630 genera 
and 10,400 species of trees, shrubs, lianas and a few 
herbs. The family is distributed worldwide, especially 
in tropical to warm termperate settings. The fossil rec- 
ord of this large family is rather poorly known, but 
there are a few unequivocal records of the family in 
the Eocene of the northern Hemisphere, including the 
taxon described below. 


Genus EMMENOPTERYS Oliv. 


Emmenopterys dilcheri sp. nov. 
Plate 36, figures 1-11; Text-figure 15 


Etymology.—This species is named for David L. 
Dilcher, recognizing the importance of his contribu- 
tions to angiosperm paleobotany. 

Description.—Cymes with pedicellate, elongate fruits; 
infructescence axis at least 20 mm long, 0.5 mm thick, 
pedicels 8-10 mm long, with bract scars subtending 
each pedicel; fruits spindle-shaped, bilocular, septici- 
dal multi-seeded capsules, fusiform in longitudinal 
view, circular to elliptical in cross section; base round- 
ed-acute, tapering smoothly to the pedicel attachment, 
without a scar at the junction of the pedicel with the 
base of the fruit; apex acute, with short epigynous са“ 
lyx; fruit length 15-20 mm, width measured perpen- 
dicular to the septum 6.2-7.0 mm, thickness in the 
plane of septum 3.6-4.0 mm; length/width ratio Dp 
2.8; exterior of fruit (exocarp?) smooth, with widely 
spaced longitudinal ribs; each carpel D-shaped in cross 
section, placentation axile, with a T-shaped placenta 
protruding nearly halfway into each locule; епдосагр 
wall thin (approx. 0.125 mm), with transversely ori- 
ented fibers adjacent to locule; septum thin, approxi- 
mately equal to thickness of endocarp wall; seeds пи- 
merous, imbricate, flattened and elongate parallel t0 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 81 


the long axis of the fruit; seed ca. 3.0 mm long, 1.1- 
1.4 mm wide, 0.3-0.4 mm thick; consisting of an el- 
liptical central body surrounded by a loosely adherent 
wing-like seed coat; seed coat with an outer layer of 
large cells 100-220 um in diameter, four- to seven- 
Sided in face view, with thick anticlinal walls but thin 
Outer periclinal walls that are frequently collapsed; these 
cells reticulately thickened to form a grid of small po- 
lygonal depressions, 25-30 um in diameter. 

Specimens.— Holotype: OMSI Pb1197. Paratypes: 
OMSI Pb1307, UF 9333, 9494-9499, USNM 435091. 

Discussion. — Emmenopterys dilcheri is represented 
by compressions of the infructescence (Pl. 36, fig. 1), 
and by casts and permineralizations of individual fruits. 
The holotype (Pl. 36, figs. 2, 3) shows the arrangement 
Of fruits in the infructescence and, upon sectioning, 
Tevealed the bilocular structure and axile placentation 
(Р]. 36, figs. 4, 5). An especially well preserved isolated 
fruit was also sectioned (Pl. 36, fig. 6) to reveal details 
of Seed position and morphology. The seeds, as seen 
In transverse (Pl. 36, figs. 7, 8) and longitudinal section 
(PI. 36, figs. 9-11) show a distinctive seed coat of large 
reticulately thickened cells. 

This species conforms to the Rubiaceae in the cy- 
Mose infructescence, inferior ovary, bilocular fruit, ax- 
lle placentation and endocarps with transversely ar- 
ranged fibers. Capsular fruits occur in several tribes of 
the family (Schumann, 1891). Reticulately thickened, 
Or “pitted” seed coats are characteristic of the subfam- 
ily Cinchonoideae (Bremekamp, 1966), and are con- 
sidered to be plesiomorphic in the Rubiaceae because 
they also occur in Loganiaceae (Bremer and Struwe, 
1992). The fossil is placed confidently in the tribe Cin- 
choneae (sensu Schumann, 1891) on the basis of ped- 
Icellate, elongate, thin-walled, septicidal capsules, with 
Numerous tiny winged seeds (Standley and Williams, 
1975). The Cinchoneae have about 40 genera occurring 
m tropical and subtropical regions. Genera with elon- 
8ate, bilocular septicidal capsules particularly similar 
to the fossil include: Badusa (New Guinea, western 

acific), Cosmibuena (tropical America), Emmenop- 
lerys (China), Exostema, Ferdinandusa (tropical 
America), Ladenbergia (tropical America) and Luculia 
Himalayas and Yunnan). These genera can be distin- 
Suished from one another by the size and extent of 
lobing of the persistent calyx and by features of seed 
Morphology including size and shape of the wing, and 
the patterns of thickening of cells in the seed coat. The 
fossil most closely resembles extant Emmenopterys, to 
Which it is assigned. Emmenopterys dilcheri compares 
Well with extant E. henryi, a deciduous tree endemic 
10 the mixed mesophytic forests of China, in the rel- 
atively inconspicuous persistent perianth, and in fea- 
tures of the seed including elongate wing shape (Pl. 36, 


fig. 12) and reticulate thickening of the seed coat cells 
(Pl. 36, fig. 13). 

Emmenopterys dilcheri is the earliest record of fruits 
of Rubiaceae from North America. The same family 
has also been recognized on the basis of cuticularly 
preserved leaves from the middle Eocene Claiborne 
Formation of western Kentucky and Tennessee (Roth 
and Dilcher, 1979). Leaves of Paleorubiaceophyllum 
Roth et Dilcher (1979) have adnate stipules similar to 
those occurring in the genera Condaminea and Cos- 
mibuena of the subfamily Cinchonoideae (tribes Con- 
damineae and Cinchononeae, respectively). Taken to- 
gether, these fruit and leaf records indicate that the 
subfamily Cinchonoideae of the Rubiaceae was well 
established in North America by the middle Eocene. 


Family SABIACEAE Blume 


The Sabiaceae are a family with three genera. Sabia, 
with about 20 species, is a genus of scandent shrubs, 
or rarely erect plants, distributed in eastern and south- 
eastern Asia, Indomalesia and the Solomon Islands. 
Meliosma, distributed in Asia and central America, 
and Ophiocaryon of Central American distribution, are 
mainly trees and shrubs. Meliosma and Sabia are well 
represented in the fossil record of the northern hemi- 
sphere. The Sabiaceae are represented in the Nut Beds 
by several species of Meliosma and one of Sabia. 


Genus MELIOSMA Blume 


Meliosma is a genus of trees and shrubs found in the 
tropics and subtropics of eastern Asia, Malesia, Central 
America and the West Indies. It is well represented by 
endocarps in the late Cretaceous (Knobloch and Mai, 
1986; Crane, Manchester and Dilcher, 1990, p. 32) 
and early Tertiary (Reid and Chandler, 1933; Mai, 
1975; Crane et al., 1990) of Europe and North Amer- 
ica. Meliosma is one of the most species-rich genera 
in the Nut Beds flora with five distinct species based 
upon fruits and seeds: M. beusekomii sp. nov., M. bo- 
nesii sp. nov., M. elongicarpa sp. nov., M. cf. jenkinsii 
Reid et Chandler, and М. leptocarpa sp. nov. In ad- 
dition, the genus is represented at the same locality by 
leaves (Manchester, 1981) and wood (Manchester, un- 
published). Leaves are reported from several other Ter- 
tiary localities in western North America (Taylor, 1990). 

Van Beusekom (1971) emphasized the importance 
of endocarp characters in distinguishing extant sub- 
genera, sections and species of Meliosma. Endocarps 
of Meliosma subgenus Kingsboroughia (Liebm.) Beus. 
are distinguished from those of subgenus Meliosma by 
the lack of a funicular canal within the endocarp. In 
subgenus Kingsboroughia, the vascular supply to the 
seed enters directly through a ventral pore in the en- 
docarp, without passing through a funicular canal. Sub- 


82 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


genus Meliosma is divided into two sections, Meliosma 
(Asian) and Loranzanea (Liebm.) Beus. (Central and 
South American), that can be differentiated by the mor- 
phology of the funicular canal morphology. In section 
Meliosma, the funicular canal is ventrally recessed 
within the endocarp, whereas in section Lorenzanea, 
the canal is ventrally extended. These differences are 
especially clear in longitudinal sections in the plane of 
symmetry as illustrated by Van Beusekom (1971, pp. 
366-367, fig. 4), and are useful in the analysis of fossil 
remains (Pl. 38, figs. 1-4). According to these char- 
acters, the Nut Beds species correspond to subgenus 
Kingsboroughia (M. cf. jenkinsii), and to Meliosma 
subgenus Meliosma, section Meliosma (M. beuseko- 
mii, M. bonesii, M. elongicarpa, M. leptocarpa). 


Subgenus MELIOSMA sensu Beus. 
Section MELIOSMA sensu Beus. 


Meliosma beusekomii sp. nov. 
Plate 37, figures 1—5; Plate 38, figure 1; 
Text-figure 16A 


Etymology.—This species is named after C. F. Van 
Beusekom, recognizing the importance of his contri- 
butions to the systematics of Meliosma. 

Description.—Endocarp subglobose, bilaterally sym- 
metrical, obliquely truncate ventrally with a prominent 
circular funicular cavity or plug, and with a pro- 
nounced keel in the plane of symmetry; unilocular, 
single-seeded; dorsiventral height 3.2-5.7, аур. 4.2 mm 
(SD=0.61, n=31), length in plane of symmetry per- 
pendicular to height 3.2-5.2, avg. 4.0 mm (SD=0.56, 
n-31), width across plane of symmetry 3.0—5.0, avg. 
3.7 mm (SD=0.49, n=31); height/width ratio 1.1; gen- 
tle ridges on the lateral faces delimiting 5 to 10 facets 
or shallow depressions on each side of the median keel; 
lower lateral surface convex-rounded; funicular canal 
passing obliquely into the locule, 1.8 mm long; en- 
docarp wall 0.4 mm thick, woody, composed of 10- 
12 rows of densely packed longitudinally elongate un- 
dulate-walled (“‘centipede-like’’) sclereids. Locule cast 
similar in shape to the endocarp, with more prominent, 
angular funicular cavity, and surface with small elon- 
gate punctae more or less aligned in vertical rows; seed 
conforming to the shape of the locule. 

Specimens.— Holotype: UF 6673. Paratypes: HU 
59978, UF 6672, 6683, 6674-6692, 6694-6705, 
USNM 435119 (lot of 60 specimens). 

Discussion.— This is the most abundant species of 
Meliosma in the Nut Beds. It is represented by per- 
mineralized endocarps (Pl. 37, figs. 1-2; Pl. 38, fig. 1), 
and by locule casts (Pl. 37, figs. 3-5). The species is 
distinguished by facets or depressions on the lateral 
faces. The morphology of the endocarp wall in the 


funicular area, as seen in longitudinal section (Pl. 38, 
fig. 1; Text-fig. 16A) conforms to the pattern that Van 
Beusekom (1971) described as diagnostic for subgenus 
Meliosma, section Meliosma, and differentiates the 
species from the extant American section Lorenzanea. 
Among extant species, it is most similar in endocarp 
morphology to M. simplicifolia (Roxb.) Walp. and M. 
pinnata (Roxb.) Walp. 


Meliosma bonesii sp. nov. 
Plate 37, figs. 6—9; Plate 38, fig. 2; 
Text-figure 16B 


Etymology.—The epithet recognizes the major con- 
tributions of Thomas J. Bones through his persistent 
collecting at the Clarno Nut Beds locality. 

Description.—Endocarp subglobose, bilaterally sym- 
metrical, obliquely truncate ventrally with a prominent 
circular funicular cavity or plug and a pronounced me- 
dian keel in the plane of symmetry; unilocular, single- 
seeded; dorsiventral height 3.8-7.6, avg. 5.5 mm 
(SD=1.34, n=7), length in plane of symmetry perpen- 
dicular to height 4.9-7.2, avg. 5.8 mm (SD=0.79, n=7), 
width across plane of symmetry 4.6—7.5, avg. 5.7 mm 
(SD=0.93, n=7); height/width ratio 0.8-1.0; lateral 
faces smooth and rounded; lower lateral surface more 
or less straight; funicular canal passing obliquely into 
the locule, 3.2 mm long; endocarp wall 0.6 mm thick, 
woody, composed of elongate sinuous-walled sclereids. 
Locule cast with more prominent, angular funicular 
cavity, and surface with small elongate punctae more 
or less aligned in vertical rows. 

Specimens.—Holotype: UF 6714. Paratypes: UF 
6712-6717, 6802, 6803, 6811-6816, 6825-6833. 

Discussion.—Meliosma bonesii is represented by 
permineralized endocarps (Pl. 37, figs. 6, 7) and locule 
casts (Pl. 37, figs. 8, 9). Endocarps of this species tend 
to be larger than those of M. beusekomii, but there is 
some overlap in size range. Morphologically, M. bo- 
nesii is distingished from M. beusekomii by smooth, 
rounded, rather than faceted lateral faces, and by the 
more or less straight rather than convex basal side. 
Longitudinal sections in the plane of symmetry show 
a well-developed oblique funicular canal (Pl. 38, fig. 
2; Text-fig. 16B) with morphology conforming to that 
indicated of subgenus Meliosma, section Meliosma 
(Van Beusekom, 1971). М. bonesii shows thicker walls 
and a more elongate locule) than М. beusekomii (cf. 
Text-figs. 16B and 16A). 


Meliosma elongicarpa sp. nov. 
Plate 37, figures 10-12; Text-figure 16D 


Etymology.—elongatus (L = prolonged) + karpos 
(Gr = fruit), referring to the elongate form of the en- 
docarp. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 83 


іст 


Text-figure 16.— Meliosma species from the Nut Beds. A, Meliosma beusekomii sp. nov., ventral view and longitudinal section in plane of 
bisymmetry showing oblique funicular canal. B, M. bonesii sp. nov., ventral view and longitudinal section in plane of bisymmetry. C, M. 
leptocarpa sp. nov. ventral view and longitudinal section in plane of bisymmetry. D, M. elongicarpa sp. nov., ventral and lateral views. E, 
M. cf. Jenkinsii Reid et Chandler, ventral view and longitudinal section in plane of symmetry. 


Description. —Endocarp elongate-ellipsoidal, bilat- 
erally symmetrical, with circular ventral funicular plug, 
and with a median keel in the plane of symmetry, 
unilocular, single-seeded; dorsiventral height 9.8 mm, 
length in plane of symmetry perpendicular to height, 
5.0 mm, width across plane of symmetry 4.4 mm, 
height/width ratio 2.2; lateral faces smooth and round- 
ed; lower lateral side convex, rounded; funicular canal 
Passing obliquely into the locule. 

Specimen.—Holotype: UCMP 10701. 

Discussion.—This species, known only from a few 
Specimens, is distinguished from the other Clarno spe- 
“les by its elongated height, which results in a height/ 
Width ratio of about 2; the other species have height/ 
Width ratios of about 1. The morphology is consistent 
With placement in section Meliosma. Larger, but sim- 
Папу elongate endocarps occur іп extant М. sumatrana 
(Jack) Walp. of Sumatra. 


Meliosma leptocarpa sp. nov. 
Plate 37, figures 16-19; Plate 38, figure 4; 
Text-figure 16C 


Etymology.— Leptos (Gr = thin, slender) + carpum 
Gr = fruit) referring to the narrow width of the en- 
docarps. 

Description.—Endocarp ellipsoidal, bilaterally sym- 
Metrical, laterally compressed in the plane of sym- 


metry; with a pore on the ventral surface, finely ru- 
gulate, with a median keel running all around the 
endocarp in the plane of symmetry; unilocular, single- 
seeded, dorsiventral height 4.0-5.1, avg. 4.6 mm 
(SD=0.36, n=17), length in plane of symmetry per- 
pendicular to height 2.8-4.5, avg. 3.4 mm (SD=0.52, 
n=17), width across plane of symmetry 2.0-2.6, avg. 
2.2 mm (SD=0.18, n=17); height/width ratio 1.8-2.4, 
surface rugose; endocarp wall 200 um thick, composed 
of longitudinally elongate ““centipede-like” cells. Loc- 
ule casts similar in shape to Ше endocarps, but smooth- 
surfaced, height 2.8-5.0, avg. 4.0 mm (SD=0.50, n=16), 
length 2.5-4.0, avg. 3.0 mm (SD=0.34, n=16), width 
1.5-2.3, avg. 2.0 mm (SD=0.17, n=16). 
Specimens.—Holotype: UF 6725. Paratypes: UF 
6739-6768, 6817-6819, USNM 355377 (8 speci- 
mens), 355430, 355435 (10 specimens), endocarps. 
Discussion.—This species is very common in the 
Nut Beds. It is distinguished by its very narrow width 
and prominent surface ruminations of the endocarp. 
At first I was hesitant to place this species in Meliosma, 
because the endocarps are more compressed in the 
plane of symmetry than any other extant or fossil spe- 
cies of the genus. However, longitudinal sections (Pl. 
38, fig. 4; Text-fig. 16C) reveal internal morphology 
and anatomy diagnostic of Meliosma and funicle as in 
other species of Meliosma. The morphology of the 
endocarp wall in the funicular area, as seen in longi- 


84 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


tudinal section, indicates that this species conforms to 
the Asian section Meliosma. 


Subgenus KINGSBOROUGHIA sensu Beus. 
Section KINGSBOROUGHIA sensu Beus. 


Meliosma cf. jenkinsii Reid et Chandler 
Plate 37, figures 13-15; Plate 38, figure 3; 
Text-figure 16E 


Description.—Endocarp large, subglobose, bilater- 
ally symmetrical, smooth, ventral side marked by a 
prominent rounded triangular funicular plug; with a 
pronounced median keel in the plane of symmetry; 
unilocular, single-seeded; dorsiventral height 8.0-13.2, 
аур. 11.4 mm (SD=1.8, п=7), length in plane of sym- 
metry perpendicular to height, 8.5-13.2, avg. 11.0 mm 
(SD=1.7, n=7), width across plane of symmetry 9.0- 
12.2, avg. 10.4 mm (SD=1.4, n=7), height/width ratio 
0.9-1.0; endocarp wall 0.8-1.0 mm thick, woody, com- 
posed of elongate undulating cells. Locule cast con- 
forming in shape to the endocarp, smooth, rounded, 
with prominent ventral cavity and pronounced median 
keel, height 9.0-11.4, avg. 10.4 mm (SD=1.2, n=7), 
length 7.4-11.5, avg. 9.9 mm (SD=1.5, n=7), width 
7.8-11.3, avg. 9.2 mm (SD=1.6, n=7). 

Specimens.—UF 6706, 6707, 6786, 9405, 9851, 
USNM 355193, 435084, 446060, 446065, 446059. 

Discussion.— Meliosma cf. jenkinsii is distinguished 
from other Clarno species of Meliosma by its larger 
size, smooth contours and more massive ventral plug. 
The funicular canal is short, and not markedly oblique. 
The species is represented by a single complete en- 
docarp and several locule casts, some with patches of 
the endocarp wall remaining. The range of endocarp 
size was extrapolated by adding the thickness of the 
endocarp wall (0.9 mm х 2) to the locule cast dimen- 
sions. 

This species corresponds in size and shape to Me- 
liosma jenkinsii Reid et Chandler from the London 
Clay (Reid and Chandler, 1933; Chandler, 1961b), and 
Geiseltal (Mai, 1975) floras. Van Beusekom (1971) 
states that the fossils “should undoubtedly be placed 
in Subgenus Kingsboroughia Section Kingboroughia by 
the fact that the vascular bundle running through the 
ventral pore of the endocarp wall is situated 1n a ventral 
groove in the latter, and is still present in the shape of 
a spatulate ventral ‘plug’ at the ventral side of the 
endocarp.” Longitudinal sections of the Clarno spec- 
imens support Van Beusekom’s conclusion (PI. 38, fig. 
2; Text-fig. 16E). Meliosma Section Kingsboroughia is 
disjunct today between central America and Asia with 
two species, M. alba and M. rufo-pilosa. 


Genus SABIA Colebr. 
Sabia prefoetida (Becker) comb. nov. 
Plate 38, figures 5-8 


Symplocarpus prefoetidus Becker, 1969, p. 10, pl. 13, figs. 19, 20. 
Sabia, Bones, 1979, pl. 3, fig. 14. 


Description.—Endocarp unilocular, single-seeded, 
laterally compressed, bilaterally symmetrical, resem- 
bling a painter’s pallete in lateral view, rounded in 
lateral outline except for a ventral notch containing the 
funicle; length (longest dimension) 3.3-5.5, avg. 4.5 
mm (SD=0.55, n=11), width across plane of symmetry 
1.1-1.6, avg. 1.4 mm (SD=0.15, n=11), height (at right 
angle to length) 3.1-4.8, avg. 4.1 mm (SD=0.40, n=1 1); 
keel in the plane of symmetry running from the basal 
scar nearly 320° around the fruit terminating in a sharp 
rostrum adjacent to the ventral notch, both lateral faces 
with about 30 shallow crater-like depressions, which 
are largest in the center, decreasing in size toward pe- 
riphery. 

Specimens.—HU 59977, 59989, UCMP 10652- 
10654, UF 5245, 6727-6736, 6801, 6822-6824, 
USNM 353968. Holotype: YPM-NYBG 894 from 
Beaverhead Basin, Montana. 

Discussion.—Sabia is represented in the Nut Beds 
by endocarp and locule casts. Although no cellular de- 
tails are preserved, the shape, size and surface sculpture 
of these casts correspond closely to extant Sabia. The 
size and sculpture correspond very closely to the en- 
docarp impressions that Becker (1969) illustrated from 
the Christensen Ranch locality of the Beaverhead basin 
of southwestern Montana. Reexamination of Becker’s 
original specimens, now at Yale University, has con- 
firmed that they are endocarp impressions having 
nothing to do with the spadices of Symplocarpus. Sabia 
has a good fossil record in Europe, ranging from the 
late Cretaceous of Czechoslovakia and Germany 
(Knobloch and Mai, 1986), to the Miocene of Poland 
(Czeczott and Skirgiello, 1959) and Germany (Gregor, 
1978) to the Pliocene of France (Geissert and Gregor, 
1981). 


Family SAPINDACEAE Juss. 


The Sapindaceae are a large family of 144 genera 
and 1325 species of tropical to temperate trees, shrubs, 
lianas and herbaceous climbers. Fruits of Sapindaceae 
range from berries to samaras (Van Welzen, 1988). 
Seeds of the family are exalbuminous, with highly char- 
acteristic embryos that completely fill the seed. Reid 
and Chandler (1933) observed that “the curved, ta- 
pering radicle, sometimes large, sometimes small, 15 
contained in a pocket of the seed coat. The large cot- 
yledons may lie upon or beside one another, and be 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 85 


Without folds; or they may lie upon one another and 
be folded or coiled together in a more or less compli- 
Cated manner.” Because the superficial characters of 
the embryo are impressed more or less distinctly on 
the inner surface ofthe seed coat, these characters may 
also be observed in fossil seeds. 

The Sapindaceae are represented in the Nut Beds by 
three taxa: Deviacer wolfei gen. et sp. nov., and two 
Species of Palaeoallophylus. Elsewhere in the Clarno 
Formation, the family is represented by impressions 
Of the distinctive bladder-like fruits of Koelreuteria 
(Sheep Rock Creek locality, Manchester, unpublished). 
The Aceraceae, which logically should be treated as a 
tribe within Sapindaceae (Radlkofer, 1890; Muller and 
Leenhouts, 1976), have not been identified among the 

ut Beds fruits and seeds, although Acer fruits occur 
at the Sheep Rock Creek locality of the Clarno For- 
Mation (Wolfe and Tanai, 1987), and fruits of Dipter- 
Ота occur at the Sheep Rock Creek, Dry Hollow, and 
White Cliffs localities. 


Genus DEVIACER gen. nov. 


Etymology.—Devius (L = turn or bend aside; devi- 
ate) + Acer (L = maple), referring to the backward 
Orientation of the fruits with respect to those of Acer. 

Type species.— Deviacer wolfei sp. nov. 


Deviacer wolfei sp. nov. 
Plate 39, figures 1-6; Text-figure 17B 


Etymology.— The epithet is in honor of Jack A. Wolfe 
Tecognizing his contributions to paleobotany and the 
Systematics of Sapindales. 

Description. —Samara bilaterally symmetrical, with 
m elongate lateral wing in the plane of symmetry ad- 
Joining an elliptical nutlet; samara length 20-34, avg. 
24 mm (SD-4.7, n-8), width 5.0-10.3, avg. 7.7 mm 
(SD=1.8, n=8); pedicel/stipe (proximal keel) 3-4 mm 
long, without obvious perianth scars; nutlet elliptical, 
length 5.0-8.0, avg. 6.2 mm (SD=1.8, n=7), width 2.2- 
4.5, avg. 3.2 mm (SD=0.82, n=7), up to about 2.5 mm 
thick with reticulate ribbing pattern over the surface, 
Tounded laterally except for a flattened abscission scar 
Оп the dorsal margin; this scar oriented at 40—60° to 
the trend of the wing; wing dorsal margin more or less 
Straight to slightly convex, rounded distally; ventral 
Margin concave proximally forming a sulcus adjacent 
to the nut, convex, rounded distally; venation subpar- 
allel, dichotomizing, running parallel to the dorsal 
Margin for some distance then arching distally toward 
the ventral margin; veins closer together and coalesced 
along the straight dorsal margin of the wing; a rudder- 
Ike Projection 1.0-1.2 mm long arises from the nut 
9Pposite the pedicel attachment and protrudes parallel 


Text-figure 17.—Comparison of Acer and Deviacer samaras. A, 
Acer grandidentatum. B, Deviacer wolfei gen. et sp. nov. 


to the long axis of the seed, with its tip recurved toward 
the wing. 

Specimens.— Holotype: OMSI Pb1133. Paratypes: 
OMSI Pb489, 567, 590, 632, 633, 634, 638, 724, 802, 
930, 1107, 1110, 1117, 1124, 1129, 1130, 1321, 1325, 
1425, 1521, 1524, 1529, 1576, 1593, 1594, 1774, 1786. 

Discussion. — Deviacer is common at the Nut Beds 
in a small area of the basal leaf layer at the north end 
of Face 3, preserved as impressions showing good de- 
tails of the venation (Pl. 39, figs. 1—5). It is also known 
from impressions at other Clarno localities including 
West Branch Creek, White Cliffs (Pl. 39, fig. 6) and 
Gosner Road. This material appears to be conspecific 
with that from the middle Eocene of British Columbia 
attributed to Acer sp. by Mathewes and Brooke (1971). 

Referring to this type of samara informally as the 
“Acer” arcticum type, Wolfe and Tanai (1987) pointed 
out that it may represent an extinct genus of Aceraceae. 
The samaras resemble those of Acer in shape of the 
seed, the coarse reticulum of veins over the nutlet, and 
in the shape and venation of the wing (Pl. 39, figs. 1— 
6), yet they differ fundamentally in mode of attach- 
ment. Whereas in Acer the attachment scar is situated 
on the ventral margin of the nutlet and forms an acute 
angle with the thickened margin of the wing (Text-fig. 
17A), in Deviacer the scar is on the dorsal margin of 
the nutlet and forms an obtuse angle with the thickened 
lateral margin of the wing (Text-fig. 17B). The large 
flat attachment scar indicates that the fruits were schiz- 
ocarpic, with two or more fruits sharing the same axis, 
as in fruits of Acer. In Acer schizocarps the ventral 
wing margins of adjoining samaras face each other. 
Although fruits of Deviacer have not been found in 
connection, the attachment angle indicates that the 
dorsal margins of the wings would have faced each 
other. As noted by Wolfe and Tanai (1987), this ori- 
entation is encountered among extant samaroid Sap- 
indaceae; however, “in samaroid Sapindaceae (pri- 
marily Paullinieae), the samaras are typically attached 


86 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


along the length ofthe wing (as well as along the nutlet) 
and the veins are not strongly coalesced along one side 
of the wing; this last character is valid also for sapin- 
daceous samaras that are attached only along the nut- 
tet 

It is not known whether Deviacer samaras were borne 
in pairs as in Aceraceae or in threes as is typical of 
Sapindaceae. Although the family name Aceraceae is 
well ingrained in the literature, there is no firm basis 
for treating it separately from the Sapindaceae. Radlk- 
ofer (1890) concluded on macromorphological grounds 
that the Aceraceae should logically be treated as a tribe 
within Sapindaceae, an interpretation that is also sup- 
ported on the basis of pollen morphology (Muller and 
Leenhouts, 1976). Deviacer is thus treated as an extinct 
genus of Sapindaceae sensu lato. 


Genus PALAEOALLOPHYLUS Reid et Chandler 


The generic name Palaeoallophylus denotes simi- 
larity in pattern of folding of the embryo to that seen 
in extant Allophylus. However, as noted by Reid and 
Chandler (1933) when they established this genus, sim- 
ilar patterns of embryo folding may occur in other 
extant genera of this large family. Although assignment 
of these fossils to the Sapindaceae is sound, other char- 
acters would be needed to determine how closely the 
fossil is allied to extant Allophylus. 


Palaeoallophylus globosa sp. nov. 
Plate 38, figures 9, 10 


Description. —Seed globose, bilaterally symmetrical, 
with a large folded and coiled embryo; length 3.6-6.0, 
avg. 4.6 mm (50-0.52, n=30), width 3.7-5.2, avg. 4.5 
mm (SD=0.45, n=29), thickness (3.2) 3.5-5.2, avg. 4.1 
mm (SD=0.55, n=29). Length/thickness ratio 0.8-1.2; 
hilar scar medial, adjacent to radical; seed coat smooth, 
thin (0.1-0.2 mm), forming a pocket around the radical 
% as long as the seed; chalaza marked by a transverse 
band inside the seed coat situated about и of the cir- 
cumference from the hilum; seed exalbuminous, em- 
bryo large, consisting of two cotyledons folded one 
inside the other like a bent knee, encircled by a coil of 
the hypocotyl and radicle; radicle tapering laterally. 

Specimens.— Holotype: UF 6587. Paratypes: HU 
59965, OMSI Pb194, UF 6585, 6586, 6589-6595, 
USNM 312766 (Bones, 1979, pl. 4, fig. 1), 355041- 
355043, 355046, 355055, 355058, 355059, 424852- 
424857, 424859—424871. 

Discussion.— Palaeoallophylus globosa is represent- 
ed by internal molds of the seed coat; usually the seed 
coat itself is not preserved except in the radicle region. 
These molds clearly show the impression made by the 
large folded and coiled embryo (Pl. 38, figs. 9, 10). The 
combination of a coiled embryo and a pocket in the 


seed coat containing the radicle is characteristic of ex- 
tant and fossil Sapindaceae as noted by Reid and Chan- 
dler (1933). This species is similar in shape, but some- 
what smaller than Palaeoallophylus rotundatus Reid et 
Chandler from the London Clay flora (Reid and Chan- 
dler, 1933). 


Palaeoallophylus gordonii sp. nov. 
Plate 38, figures 11-16 


Etymology.— This species is named for Ian Gordon, 
who provided extensive help with paleobotanical field 
work in the Clarno Formation. 

Description.—Seed elongate ovoid, bilaterally sym- 
metrical, rounded apically and truncate to obtuse-con- 
ical at the hilar end; length 4.8—6.0, avg. 5.4 mm 
(SD=0.44, n=13), width 3.3-4.9, avg. 4.1 mm 
(SD=0.43, n=13), thickness 3.2-4.5, avg. 3.6 mm: 
(SD=0.43, n=13), length/thickness ratio 1.3-1.8; seed 
coat 75-100 um thick, smooth, forming a pocket around 
the radicle about % of the length of the seed; seed 
exalbuminous, embryo large, consisting of two coty- 
ledons folded one inside the other in an *S" configu- 
ration; radicle tapering toward the hilum. 

Specimens.— Holotype: USNM 424669. Paratypes: 
UCMP 10730, 10731, UF 6584, 6596, 6597, USNM 
355048, 355051 (10 specimens), 355060, 424666- 
424668, 424670. 

Discussion.— This species is represented by internal 
molds of the seed coat, showing the impression of the 
folded cotyledons and the protruding radicle (Pl. 38, 
figs. 11, 12), sometimes with remains of the seed coat 
adhering (Pl. 38, figs. 13-16). The seeds of this species 
are narrower, more elongate, than those of P. globosa. 
Itis similar in morphology, but somewhat smaller than 
Palaeoallophylus minimus Chandler (1961b; = Sap- 
indospermum sp. 4, Reid and Chandler, 1933) from 
the London Clay. 


Family SAPOTACEAE Juss. 


The Sapotaceae are a family of about 107 extant 
genera and 1000 species of trees and shrubs, mostly 
tropical in distribution. The seeds described below 
confirm the presence of this family in the Nut Beds. 


Genus BUMELIA Sw.? 


Bumelia? globosa sp. nov. 
Plate 39, figures 7-16 


Description.—Seed anatropous, subglobose; internal 
mold of seed coat bilaterally symmetrical, circular in 
cross section, slightly longer than wide, with an oblique, 
apical truncation, base rounded to slightly pointed; 
length 3.3-5.9, avg. 4.5 mm (SD=0.49, n=25), equa- 
torial diamter 3.2-4.4, avg. 3.9 mm (SD=0.25, n=25); 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 87 


truncate face flat, circular in outline, perpendicular to 
the plane ofsymmetry, sloping toward the front ofseed 
at 70° to the long axis of the seed; hilum a small scar 
near the center of Ше truncation, slightly offset toward 
the edge along which the raphe travels; micropyle ev- 
ident as a tiny pore at the highest end of the truncation; 
raphe bundle passing in the plane of symmetry from 
the hilum across the front margin of seed to the chalaza 
3t the base; with a network of veins radiating from the 
hilum over the truncate surface. 

Specimens.— Holotype: USNM 424644. Paratypes: 
HU 60000, UCMP 10645-10649, UF 8668, 8669, 
8672-8685, USNM 353964 (21 specimens), 353965 
(4 specimens), 355446, 355488 (6 specimens), 424656- 
424664, 

Discussion. — This species is represented by trans- 
lucent chalcedony internal molds ofthe seed coat. The 
Taphe is sometimes preserved as an opaque band that 
Can be observed in silhouette within the translucent 
chalcedony (PI. 39, fig. 10). The seed coat itself is not 
Preserved. 

The morphology of these specimens closely matches 
that of the cavity inside the hard seed coat of extant 
Bumelia lacuum. They correspond in size, bisymme- 
try, angle of the truncation, the nearly hemispherical, 
but subtly pointed base, relative spacing of the hilum 
and micropyle, course and thickness of the raphe. Be- 
Cause the Nut Bed fossils are missing the seed coat, 
the resemblance of these fossils to seeds of extant Bu- 
Melia is not immediately obvious. However, if a mold 
were to be made of the inside of the seed coat of an 
extant seed, it would be difficult to distinguish from 
these fossils. The assignment to Bumelia? includes a 
Question mark because my study of extant seeds in this 
family has not been sufficiently thorough to establish 
that the morphology of these seeds is unique to that 
genus. 


Bumelia? subangularis sp. nov. 
Plate 39, figures 17-21 


Etymology.—The epithet refers to the subangular 
external morphology of the seed. 

Description.—Seed subellipsoidal, bilaterally sym- 
Metrical, smooth, obliquely truncate apically, rounded 
basally, rounded-subtriangular outline in cross section; 
length 6.0-6.9, avg. 6.6 mm (SD=0.43, n=5), width 
Across plane of symmetry 3.7-5.0, avg. 4.6 mm 
(50-0.54, n=5), width parallel to plane of symmetry 
4.0-5.4, avg. 4.6 mm (SD=0.55, n=5); truncation more 
Or less flat, obovate in outline, sloping toward front of 
Seed at 50—60° to the long axis of the seed, circular 

Паг scar in the center of the truncation; network of 
Veins radiating from the hilum in plane of truncation. 


Specimens.—Holotype: UF 8743. Paratypes: UF 
8744, USNM 424681, 435082, 435083. 

Discussion. — This species is represented by chalced- 
ony seed casts without internal structure preserved. 
The smooth, shiny surface of the fossils suggests that 
the seeds were hard and shiny, possibly from a bird- 
dispersed berry. The slight angling (Pl. 39, figs. 17, 18) 
suggests that more than one seed may have been pres- 
ent in each fruit. Because of the subangular morphol- 
ogy and larger size, Bumelia? subangularis is inter- 
preted to be a different species from the globose internal 
molds of seed coat described above as B.? globosa. 


Family SCHISANDRACEAE BI. 


The Schisandraceae contains two extant genera of 
tropical to warm temperate climbing shrubs: Schis- 
andra and Kadsura. Schisandra is distributed in Asia 
(24 species) and eastern North America (one species); 
Kadsura (22 species) is distributed from India to Ma- 
lesia. The two genera are similar in most vegetative 
and reproductive characters but are distinguished on 
the basis of a simple difference in infructescence mor- 
phology (Smith, 1947). The berries of Kadsura are borne 
together in a globose head, whereas those of Schisandra 
occur separated on an elongate spike. Seeds of both 
genera are campylotropous and shallowly c-shaped, 
with a thin, hard seed coat, cellular endosperm and 
small embryo. 


Genus SCHISANDRA Michx. 


Schisandra oregonensis sp. nov. 
Plate 40, figures 1-11 


Schisandraceae, Bones, 1979, pl. 1, fig. 8. 


Etymology.—The epithet refers to Oregon. 

Description.—Seed reniform to cashew-shaped, 
curved in plane of bisymmetry, rounded at both ends; 
length 3.7-9.9, avg. 5.6 mm (SD=1.9, n=22), dorsi- 
ventral width 3.0-7.5, avg. 4.5 mm (SD=1.4, n=22), 
thickness across plane of symmetry 1.9-5.0, avg. 2.9 
mm (SD=0.88, n=22); surface smooth; hilum situated 
in the ventral sinus; raphe originating at the hilum and 
running a medial course along the dorsal curve, di- 
minishing in prominence toward the micropylar end; 
outer surface (rarely preserved) with small craters; in- 
ner layer of seed coat smooth. Seed coat composed of: 
1) an inner uni- to biseriate layer of mostly isodia- 
metric cells 20-30 wm in diameter, 2) a prominent 
uniseriate layer of anticlinally oriented thin-walled co- 
lumnar cells 100-130 um high, 40-65 um wide, 3) a 
3- to 5-seriate layer of polygonal cells, sometimes per- 
iclinally elongate, 50-150 um wide and 30-45 um high, 
which proliferates to form a thick layer of isodiametric 
cells in the hilar sinus. 


88 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Specimens.— Holotype: USNM 424655. Paratypes: 
OMSI Pb1855, UCMP 10630, UF 6576, 6577, 6579— 
6583, USNM 312771 (Bones 1979, pl. 1, fig. 8), 355061 
(6 specimens), 355654 (3 specimens), 355061 (6 spec- 
imens), 424654 (3 specimens), 424732, 424809. 

Discussion.— Schisandra is represented by chalced- 
ony casts, molds and permineralized specimens. Con- 
spicuous features of these seeds are the curvature, dor- 
sal raphe (Pl. 40, figs. 5, 7), and the thick uniseriate 
columnar layer of the inner seed coat (Pl. 40, figs. 9- 
ђе 

Seeds of Shisandraceae аге bitegmic (Кар! and Ја- 
lan, 1964; Corner, 1976) and the inner part ofthe seed 
including the tegmen (inner seed coat) may become 
loose within the testa (outer seed coat) at maturity (Pl. 
40, figs. 14-18). Most of the fossil specimens are molds 
or permineralizations of the inner part of the seed, 
including the tegmen but excluding part of the testa. 
These have a smooth surface and a well developed 
hilar sinus and evidently were formed as infillings of 
the testa. External seed morphology is revealed in a 
rare chalcedony cast of a complete seed with the testa 
intact (Pl. 40, fig. 1), showing a rough, minutely cra- 
tered external surface, and a thick, lip-like rim along 
the hilar sinus. 

These seeds resemble those of both Kadsura and 
Schisandra in size, reniform shape, dorsal raphe, and 
seed coat anatomy. The uniseriate columnar layer of 
the tegmen, and the proliferation of isodiametric testa 
cells in the hilar region observed in the fossil specimens 
are similar to those of extant seeds. The seed surface 
is smooth in all extant species of Kadsura and in some 
extant species of Schisandra (Pl. 40, fig. 12), but rough- 
surfaced seeds occur only in Schisandra (Pl. 40, fig. 
13). The latter feature provides support for assignment 
of the fossil to Schisandra rather than Kadsura. The 
seed size of this Schisandra oregonensis is variable, 
such that the largest specimens are about twice as big 
as the smallest ones. Dimensions may have varied with 
the number of seeds per berry; there may be from one 
to five seeds in berries of extant species. 

Although Schisandra has one extant species in east- 
ern North America, the Clarno occurrence is the only 
known North American fossil record. The genus has 
also been’ determined on the basis of cuticularly pre- 
served leaves from the Eocene Geiseltal flora of Ger- 
many (Jahnichen, 1976) and on the basis of seeds from 
the Pliocene of Alsace, France (Gregor, 1981). 


Family STAPHYLEACEAE Lindley 


The Staphyleaceae are a family of five genera and 
about 50-60 species of trees and shrubs, of tropical to 
temperate distribution in the northern hemisphere. The 
family is divided into two subfamilies: Staphyleoideae 


(Euscaphis, Staphylea, Turpinea) and Tapiscioideae 
(Tapiscia, Huertea). Both subfamilies are well repre- 
sented in the Tertiary of Europe and North America 
(Mai, 1976; Tiffney, 1979; Manchester, 1988). The 
family is represented in the Nut Beds by fruits and 
seeds of Tapiscia. 


Genus TAPISCIA Oliver 


Tapiscia occidentalis Manchester 1988 
Plate 41, figures 1-9 


Tapiscia occidentalis Manchester, 1988, p. 59-66, figs. 1-11, 18, 20. 


Description.—Fruit subglobose, rounded apically, 
tapered basally to a point with a circular pedicel scar 
0.4-0.5 mm in diameter; fruit length 5.0-5.8, avg. 5.4 
mm (SD=0.40, n=6), width 4.0-4.5, avg. 4.2 mm 
(SD=0.20, n=6), thickness 3.2-4.3, avg. 3.7 mm 
(SD=0.40, n=6), pericarp thin (<0.2 mm) following 
the form of the anatropous seed; endocarp surface with 
small, regularly spaced scabrae; internal cast of seed- 
coat subglobose, length 3.7—5.4, avg. 4.6 mm (SD=1.12, 
n=18), width 3.0-4.5, avg. 3.7 mm (SD=0.40, n=18), 
thickness 2.5-4.1, avg. 3.2 mm (SD=0.47, n=8); bi- 
symetrical, rounded dorsally, pointed at the micro- 
pylar end, and concave ventrally with a prominent 
circular chalazal scar 1.5-2.5 mm in diameter, surface 
smooth to finely striate with striae running longitudi- 
nally on the dorsal surface and radiating all directions 
from the chalaza on the ventral surface; seed coat 180 
um thick, composed of five to six layers of anticlinally 
aligned elongate cells 12.5-20 um wide, 25-90 um high. 

Specimens.— Holotype: UF 5200, cast ofa fruit with 
outer layer partially broken away revealing internal 
seed cast. Additional specimens: UF 5201, 5202, 6511, 
9283, 9284, 9692, 9718-9720, USNM 354498, com- 
plete fruits; HU 59975, 59927, UCMP 10678-10686, 
UF 5203-5220, 9721 (20 specimens), USNM 312767 
(Bones, 1979, pl. 6, fig. 2), 355449-355451, 355453, 
355454 (9 specimens), 355455 (8 specimens), 355457 
(6 specimens), 435004 (Bones, 1979, pl. 6, fig. 1), seed 
casts. 

Discussion.— Tapiscia occidentalis was originally de- 
scribed from fruit and seed casts without cellular detail 
(Manchester, 1988). Some permineralized specimens 
have since been recovered (Pl. 41, figs. 4, 7-9), pro- 
viding the anatomical details included in the above 
description, and confirming the assignment to Tapis- 
cia. The species was identified through comparison 
with fruits and seeds of extant Tapiscioideae, i.e., Tap- 
iscia sinensis Oliver, Huertea cubensis Griseb., and Н. 
granadina Cuatrec. The fossil conforms to Tapiscia in 
its small size, uneven tegmen thickness, regularly spaced 
bumps on the outer surface, and the typically acute 
micropyle. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 89 


Fossil seeds of Tapiscioideae also occur in the Eo- 
Сепе Geiseltal flora of Germany (Mai, 1976). From the 
London Clay flora, Mai recognized three species of 
Tapiscia previously designated Carpolithes pusillus 
(Reid et Chandler) Chandler (1961b). Mai (1980) rec- 
ognized a fourth species from the middle Eocene Gei- 
зена! flora of Germany and provided a key for the 
identification of the five European Тарїзсїа species us- 
Ing size, shape (elongate vs. rounded), ornamentation, 
and prominence of micropyle as diagnostic characters. 
Tapiscia occidentalis is very close in morphology to T. 
Pusilla from the London Clay. Both species share the 
Same shape and external ornamentation with both hav- 
Ing a pointed micropyle. The only difference seen be- 
tween the two species is slightly larger size of T. oc- 
Cidentalis specimens. 

Tapiscia has a single living species that is a small 
tree up to 10 m high growing in the mixed mesophytic 
forests of the Yangtze-Kiang valley between Yunnan, 
western Szechuan, Hupeh and northern Chekiang in 
China (Wang, 1961). It is also recorded from wet for- 
ests to 1600 m in Kwangtung and Tonkin, China 
(Krause, 1942). 


Family SYMPLOCACEAE Desf. 


The Symplocaceae are a family of one genus, Sym- 
Plocos, with about 250 species of trees and shrubs dis- 
tributed in new world and old world tropics excluding 
Africa, and extending into temperate North America. 
Old world Symplocaceae have been monographed with 
attention to fruits and other characters by Nooteboom 
(1975). Those of the Antilles have been revised with 
attention to many ofthe same characters by Mai (1986). 
Nooteboom (1975) considered the morphology of fruits 
and of endocarps to be taxonomically useful for groups 
of Species, but refrained from making a key to indi- 
Vidual species based solely on fruit characters. Sym- 
Plocos has an excellent fossil fruit record in Europe, 
extending from Ше Eocene (Reid and Chandler, 1933; 
Chandler, 1961b) to the late Pliocene (Kirchheimer, 
1957), as reviewed in detail by Mai (1970). The fol- 
lowing constitutes the first fossil fruit record of the 
genus described from North America and is comple- 
Mentary to previous records based upon fossil pollen 
(Taylor, 1990). Symplocos fruits are also known on the 
asis of an underscribed species from the Brandon 
lignite of Vermont (Tiffney, pers. comm., 1992). 


Genus SYMPLOCOS Jacq. 


Symplocos nooteboomii sp. nov. 
Plate 41, figures 10-17 


Etymology.— This species is named in recognition 


9f the important studies of extant Symplocos carried 
Ош by Н. P. Nooteboom. 


Description.— Endocarp ovoid or ellipsoidal, some- 
what compressed laterally, slightly flared and trans- 
versely truncate apically, rounded or obtusely pointed 
basally; length 8.6-15.2, avg. 11.3 mm (SD=3.48, п=3), 
width 6.1-11.2, avg. 8.3 mm (SD=2.14, n=4), thick- 
ness 5.4—6.9, avg. 6.1 mm (50-0.67, n=4); with 13- 
14 unbranched meridional grooves or ribs (depending 
on preservation) radiating from the base and extending 
to the apical truncation; four or five locules indicated 
by distinct pores in the apical truncation. 

Specimens.— Holotype: USNM 354596. Paratypes: 
USNM 354592, 354602, 354603. 

Discussion.— Only a few specimens of Symplocos are 
known from the Nut Beds. They are preserved as chal- 
cedony casts, without anatomical preservation. En- 
docarps of this genus are so distinctive morphologi- 
cally, however, that cellular details are not required 
for identification. Assignment to Symplocos is based 
on the combination of meridional grooves, ovoid shape 
(Pl. 41, figs. 10, 11), and the truncate apex with open- 
ings corresponding to each of the locules (Pl. 41, figs. 
12, 13, 15). The apical truncation apparently corre- 
sponds to the base of the calyx as in the extant genus. 
Extant species of Symplocos have fruits that range in 
length from a few millimeters to about four centimeters 
and may have from one to five locules (Nooteboom, 
1975). 


Family THEACEAE D. Don 


The Theaceae, or tea family, is a family of about 28 
genera and 520 species of trees, shrubs and occasional 
lianas, chiefly of tropical distribution but with a few 
warm temperate representatives. The family is known 
from well-preserved fruits in the Eocene of Tennessee 
and Kentucky (Grote, 1989; Grote and Dilcher, 1989) 
and in the Tertiary of Europe (Mai, 1971). The fossil 
record of Theaceae in the northern hemisphere is re- 
viewed by Grote and Dilcher (1989). Cleyera is the 
only genus of this family that has been confirmed among 
the Nut Beds seeds. 


Genus CLEYERA Thunb. 


Cleyera grotei sp. nov. 
Plate 42, figures 1-5 


Etymology.—This species is named after Paul J. 
Grote, recognizing his contributions to the fossil his- 
tory of the Theaceae. 

Description.—Seed campylotropous, laterally flat- 
tened, rounded in outline, truncate at micropyle and 
chalaza end, with a medial condylar depression on both 
faces of the seed, around which the c-shaped embryo 
cavity curves; length 1.5-2.3, avg. 1.8 mm (SD=0.22, 
n=14), width 1.4-2.0, avg. 1.8 mm (SD=0.15, n=14), 


90 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


thickness 0.6-0.9, avg. 0.7 mm (SD=0.08, n=14); out- 
er surface reticulate with thick walled cells 100-200 
um in diameter arranged in more ог less concentric 
rows, these cells more elongate over the condyle area; 
anticlinal walls of testa cells vertical so that the lumina 
are not funnel shaped; raphe passing obliquely through 
the condyle between limbs of the embryo cavity; limbs 
of embryo cavity circular in cross section, one slightly 
longer than the other; embryo cavity cast with a seam 
all along the margin representing to the plane of de- 
hiscence. 

Specimens.— Holotype: UF 6501, seed with intact 
seed coat. Paratypes: UF 9163,9165,9449, 9450, 9529— 
9542,9543 (14 specimens), internal molds of seed coat. 

Discussion.— Cleyera grotei is represented by nu- 
merous embryo cavity casts (1.е., internal molds of seed 
coat) from the Nut Beds showing the strongly curved 
embryo cavity (e.g., Pl. 42, figs. 3, 4). The holotype is 
a rare specimen with part of the reticulate seed coat 
adhering (Pl. 42, figs. 1, 2, 5). Embryo cavity casts of 
this species closely match those from the British Ter- 
tiary that were formerly ascribed to the fossil genus 
Myrtospermum Chandler. Although Chandler (1957) 
originally suggested that the seeds were myrtaceous, 
she subsequently recognized their affinities within the 
Theaceae (Chandler, 1960, 19612), and later (Chan- 
dler, 1963b, p. 109) placed M. variabile (the type spe- 
cies of Myrtospermum) in synonymy with Cleyera (?) 
stigmosa (Ludwig) Chandler. She then transferred the 
other species formerly assigned to Myrtospermum to 
“Cleyera?” (Chandler, 1964, p. 40). 

In shape and reticulate surface sculpture of the seed, 
the Nut Beds species resembles seeds of extant Cleyera 
and Eurya of the theaceous subfamily Ternstroemo- 
ideae. Friis (1985) studied these and related extant 
genera in reviewing fossil seeds of Theaceae from the 
Miocene of Europe. Seeds of Cleyera tend to be more 
strongly campylotropous and usually larger than those 
of the other genera. Friis (1985) also observed an im- 
portant difference in the anticlinal walls of the testa 
cells that form the reticulate surface of the seed. In 
Eurya, the anticlinal walls decrease in thickness out- 
ward, so that the cell lumina are funnel-shaped, where- 
as in Cleyera, the same walls are uniformly thick so 
that the lumina are not funnel shaped (Friis, 1985). By 
this criterion, and on the basis of size and curvature, 
the Nut Beds seeds conform to Cleyera. 

Cleyera is a genus of about 17 species with a disjunct 
distribution between southeast Asia (Himalayas to Ja- 
pan) and tropical America (Mexico to Panama and 
West Indies). This genus has an excellent fossil seed 
record extending from the Eocene to the Late Tertiary 
of Europe (Mai, 1971). 


Family ULMACEAE Mirbel. 


The Ulmaceae, or elm family, includes 15 extant 
genera of trees and shrubs distributed in both the 
northern and southern hemispheres. Two subfamilies 
are recognized on the basis of differences in fruit, fo- 
liage and pollen characters (summarized in Manches- 
ter, 1989c): the Ulmoideae and the Celtidoideae. Al- 
though evidence has accumulated for the recognition 
of Celtidoideae as a separate family (Grudzinskaya, 
1967; Chernik, 1980; Takaso and Tobe, 1990), I have 
retained the traditional treatment here. The Ulmaceae 
have an excellent fossil record that includes both extant 
and extinct genera (Manchester, 1989a, c). The Nut 
Beds flora includes at least three representatives of the 
Celtidoideae (Aphananthe, Celtis, Trema) and one of 
the Ulmoideae (Cedrelospermum). Ulmus is known 
from other localities of the Clarno Formation (West . 
Branch Creek, White Cliffs, Sheep Rock Creek) on the 
basis of fruits and foliage (Manchester, 1989c) but has 
not been observed in the Nut Beds assemblage. 


Subfamily CELTIDOIDEAE 
Genus APHANANTHE Planchon 


Aphananthe maii sp. nov. 
Plate 42, figures 6-15 


Aphananthe sp. Manchester, 1989c, p. 237, fig. 12.7, I-O. 


Etymology.— This species is named for Dieter H. 
Mai, recognizing the importance of his contributions 
to paleocarpology. 

Description. — Endocarp subglobose, unilocular, sin- 
gle-seeded, pyriform in cross section, elliptical to пеаг- 
ly circular in face view; asymmetrical, length 6.5-9.0, 
avg. 7.8 mm (SD=1.05, n=5), width 5.0-6.7, avg. 6.2 
mm (SD=0.75, n=7), thickness 3.4—5.1, avg. 4.3 mm 
(SD=0.73, n—7), base rounded, apex rounded to acute, 
surface smooth to papillate; placental plug at the apex 
elliptical, oblique; endocarp with a prominent longl- 
tudinal crest deflected in course as it passes over the 
apex; endocarp wall 350—500 um thick, consisting of 
a thin papillate locule lining of more or less isodia- 
metric cells, a thick uniseriate layer of anticlinally or 
ented columnar cells 230-340 um high, 17-28 um wide, 
these cells thick-walled, with lumina closed except near 
the locule; conspicuous pits radiate from the lumina 
ofthese cells toward the locule (sometimes with crystal- 
bearing idioblasts in this layer); outside is a layer of 
translucent more-or-less rectangular cells that form ра- 
pillae at the surface, 80-150 um high, 130-180 um 
wide. Locule cast smooth, similar in shape to the en” 
docarp, but with sharper longitudinal crest. Seed with 
large embryo with a prominent coiled cotyledon. 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER oF 


Specimens.— Holotype: USNM 355693. Paratypes: 
UF 6518, 6519, 8808, 9328-9330, USNM 422384, 
424629, 424718. 

Discussion.—Aphananthe maii is represented by 
permineralized endocarps and locule casts that have 
been studied both by SEM of fractured surfaces (Pl. 
42, figs. 11, 12) and light microscopy of serial sections 
(Pl. 42, figs. 13-15). The endocarps are virtually iden- 
tical, anatomically as well as in size and morphology, 
to those of extant Aphananthe. Shared characters in- 
clude the asymmetrical longitudinal keel, the endocarp 
Wall with its thick uniseriate layer of anticlinally elon- 
gate sclereids (Pl. 42, figs. 14, 15), an outer silica layer 
forming a papillate surface (Pl. 42, figs. 6, 7), and a 
Prominently coiled embryo (Pl. 42, fig. 13). 

Aphananthe includes deciduous to semi-deciduous 
Shrubs and trees distributed today from India to China, 
Korea, Japan, Indochina, Thailand, Malesia, the Phil- 
Ippines, Java, Madagascar, eastern Australia, the So- 
lomon Islands, and Mexico. The Mexican species has 
Sometimes been placed in a separate genus, Miran- 
daceltis A.J. Sharp, but its endocarp is virtually iden- 
tical to that of the other species. This fossil species was 
illustrated but not named in a broader treatment of the 
Ulmaceae (Manchester, 1989c). Although this 1s the 
only known fossil occurrence of Aphananthe in North 
America, the genus is well-documented by endocarps 
from the Oligocene of western Siberia (Dorofeev, 1982). 


Genus CELTIS L. 


Celtis burnhamae sp. nov. 
Plate 42, figures 16—25 


Celtis sp. Manchester, 1989c, p. 240, fig. 12.8 A-C. 


Etymology.— This species is named for Robyn J. 
urnham, recognizing her contributions to the fossil 
history of Ulmaceae. 

Description. — Endocarp subglobose, bilaterally sym- 
Metrical, unilocular, single-seeded, base rounded, apex 
Tounded with a wedge-shaped tip, length 4.2-5.7, avg. 
4.8 mm (SD=0.38, n=16), width in plane of symmetry 
3.6-5.0, avg. 4.3 mm (SD=0.34, n=17), width at right 
angle to plane of symmetry 3.5-4.7, avg. 4.0 mm 
(SD=0.35, n=17); surface reticulate, with a pair of 
Strong primary longitudinal ribs forming a keel in the 
Plane of symmetry and usually a second pair of lon- 
Situdinal ribs in the plane at right angles; secondary 
rib well-developed basally, weakening apically; a net- 
Work of fine, vein-like ridges interspersed between the 
Primary and secondary ribs, delimiting areoles 0.6-0.8 
Mm in diameter; apical prominence pierced by a small 
Median funicular canal; pericarp 350-400 um thick, 
Consisting of three main layers: 1) innermost layer uni- 


seriate consisting of large cuboidal, relatively thick- 
walled cells 50-70 um in diameter, occasionally with 
rhomboidal crystals, overlain by 2) a thin uniseriate 
layer of interlocking digitate cells best seen in perider- 
mal section; combined thickness of these two layers, 
1.е., the endocarp, ca. 100 um, and 3) the outer layer, 
ca. 250—300 um thick, composed of clear silica with 
poorly preserved, apparently isodiametric cells, ca. 20 
um in diameter. Locule cast taking form of the seed 
with large curved embryo, having the same plane of 
bisymmetry as the external fruit morphology, radicle 
projecting within the apical prominence of the endo- 
carp; chalaza circular, on the opposite side of the locule 
cast from the radicle. 

Specimens.— Holotype: UF 9600. Paratypes: UF 
6520-6527, 8619-8640, 9579-9581, USNM 355466, 
424644, 435050-435052, complete endocarps; USNM 
355429, 422733, locule casts. 

Discussion.— Celtis burnhamae endocarps and loc- 
ule casts are especially abundant in the Nut Beds at 
the top of Face 3, although they are not known from 
other Clarno localities. Some specimens clearly show 
the external endocarp morphology with the character- 
istic pattern of ridges (Pl. 42, fig. 16-19); however, it 
is more common to find specimens that have broken 
along the layer of mechanical tissue giving a smooth, 
shiny appearance so that affinities to Celtis are not 
immediately obvious. The layer of interlocking digitate 
cells is seen in a few specimens that have broken per- 
idermally at the appropriate level, seen with reflected 
light microscopy (Pl. 42, fig. 25). Locule casts show the 
seed morphology with the circular chalaza at one end 
(Pl. 42, fig. 20), and the tip of the radicle at the other 
(Pl. 42, fig. 21). 

Although Celtis has a rich fossil record based upon 
fossil endocarps from the Tertiary of North America 
and Europe (Manchester, 1989c), the anatomical struc- 
ture of these fossils in relation to the modern genus 
has not been considered. Sections of some of the Clarno 
specimens (Pl. 42, figs. 22-25) reveal anatomy that 
clearly conforms to that of the extant genus, which has 
been illustrated by Chernik (1980). These endocarps 
are larger, but morphologically similar to those of С. 
edwardsii Chandler from the late Eocene of England 
(Chandler, 1963a). A clear basis for distinguishing 
among the various extant species and among different 
fossil species based upon endocarp morphology has 
not been established. It is noteworthy, however, that 
C. burnhamae the uniseriate layer of cuboidal cells 
lining the locule is three to five times thicker than the 
corresponding layer in extant species that I have stud- 
ied (C. laevigata Willd., C. occidentalis L., C. pallida 
Torr., C. sinensis Persoon). 


92 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Celtis is a genus of trees and shrubs that is wide- 
ranging in its present distribution, occurring in tem- 
perate and tropical regions of both the northern and 
southern hemispheres. 


Celtis sp. 
Plate 43, figures 1-5 


Description.— Endocarp subglobose, bilaterally sym- 
metrical, with a keel in the plane of symmetry, uni- 
locular, single-seeded, base rounded, apex pointed; 
length 3.7-4.7, avg. 4.3 mm (SD=0.30, n=20), width 
in plane of symmetry 3.0-4.1, avg. 3.7 mm (SD=0.33, 
n=20), width at right angle to plane of symmetry 2.5- 
3.3, avg. 3.0 mm (SD=0.21, n=20); apical prominence 
pierced by a small median funicular canal, surface ver- 
rucate but without ridges, except for the longitudinal 
keel. Locule cast taking form of the seed, with a large 
curved embryo, radicle protruding within the apex of 
the endocarp; pericarp ca 200 um thick with an inner 
layer of anticlinally elongate columnar cells 100—150 
um high, 50-90 um wide; surrounded in some speci- 
mens by a translucent silica layer ca. 50 um thick. 

Specimens.—UF 8640-8658, 8659-8671, 9771- 
9775. 

Discussion.— Endocarps of this species are common 
in the Nut Beds, represented by permineralized fruits 
and locule casts. The endocarps are similar in size and 
external morphology to those of Celtis burnhamae, but 
do not have a secondary ridge at right angles to the 
plane of bisymmetry, and havea verrucate (Pl. 43, figs. 
1, 2) rather than reticulate (Pl. 42, fig. 17) surface. The 
fruits are somewhat compressed in the plane of sym- 
metry (elliptical in cross section) rather than globose 
(circular in cross section) as in С. burnhamae. The 
curved embryo is apparent in specimens where part of 
the endocarp has flaked away revealing the curved loc- 
ule cast (Pl. 43, fig. 3). When sectioned transversely, 
the partition between the radicle and the rest of the 
embryo is seen as a thin line bisecting the circle of the 
locule (Pl. 43, figs. 4, 5). The wall of these fruits is thin 
relative to that of Celtis burnhamae and extant species 
that Гуе examined, but the uniseriate columnar layer 
of the endocarp compares favorably between these taxa. 

Among extant genera of Celtoideae, this species is 
most similar to Celtis. Because of the thin wall and 
lack of reticulate sculpture, it is possible that this spe- 
cies belongs in an extinct genus of the subfamily, but 
a more thorough morphological and anatomical in- 
vestigation of extant Celtis fruits is needed before an 
informed decision can be reached. 


Genus TREMA Loureiro 


Trema nucilecta sp. nov. 
Plate 43, figures 6-15 


Etymology.— Nucis (L = nut) + lectus (Г. = bed), 
referring to the Nut Beds locality. 

Description.—Endocarp subglobose, rounded-tri- 
angular in face view, bilaterally symmetrical, with a 
keel in the plane of symmetry, unilocular, single-seed- 
ed, length 2.0-3.0, avg. 2.5 mm (SD=0.21, n=16), width 
in plane of symmetry 1.9-2.6, avg. 2.3 mm (SD=0.26, 
n=15), width at right angles to plane of symmetry 1.8- 
2.4, avg. 2.1 mm (SD=0.18, n=15), base rounded, apex 
pointed with a circular scar at the tip; surface smooth; 
wall composed of uniseriate layer of anticlinally ori- 
ented columnar cells 100-120 um high and 28-60 um 
wide, these cells with fenestrate walls having numerous 
orbicular pits; apical funicle bulge composed of small 
isodiametric cells ca. 8 um in diameter; poorly pre- 
served silica layer 50 um thick surrounding Ше me- 
chanical layer. Locule cast shaped like Ше епдосагр, 
but with a sharper keel, embryo large, coiled with a 
prominent curved radicle; seed coat ca. 10 um thick, 
composed of isodiametric cells. 

Specimens.— Holotype: UF 9781. Paratypes: UCMP 
10732-10737, UF 6791-6799, 9585, 9776-9780, 
9782-9788, 9791, USNM 355097 (12 specimens), 
424635, 424723. 

Discussion. — Trema nucilecta is represented by per- 
mineralized endocarps (Pl. 43, figs. 6-10) and chal- 
cedony locule casts (Pl. 43, figs. 11-13). The locule 
casts are curved, reflecting the embryo shape (Pl. 43, 
figs. 8, 13). The specimens are similar in overall shape 
of endocarp and locule cast to Celtis sp. (see above); 
but are readily distinguished by smaller size, smooth 
exterior, and thinner, endocarp wall composed of more 
elongate columnar cells. This species conforms very 
well in size, shape, wall anatomy and embryo curvature 
to extant Trema. One difference noted between the 
fossil species and the extant species examined was the 
lack in the fossil of a secondary ridge running at right 
angles to the plane of symmetry. 


Subfamily ULMOIDEAE 
Genus CEDRELOSPERMUM Saporta 


Cedrelospermum lineatum 
(Lesquereux) Manchester 
Plate 43, figures 16-18 
Cedrelospermum lineatum (Lesquereux) Manchester, 1987b, р. 124. 


Banksites lineatus Lesquereux, 1883, p. 165, pl. 32, fig. 21. [more 
complete synoymy in Manchester 1987b]. 


Description.—Fruit a samara, consisting of a flat- 
tened elliptical endocarp adjoined by an elongate lat- 
eral wing; samara length 12.5-15.5, avg. 13.8 mm 
(SD=1.31, n=8); endocarp length 4.7—5.0, avg. 4.7 mm 
(SD=0.14, n=5), width 2.3-3.0, avg. 2.7 mm (SD=0.27, 
n=5); primary wing length 9.7-11.5 mm, width 4.5- 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 93 


5.0 mm, oriented oblique to the long axis of the en- 
docarp with 6-8 subparallel veins, including marginal 
veins that converge toward stigmatic area at the adaxial 
margin of the distal end of the wing; a small secondary 
wing 1.0-2.0 mm long, oriented nearly parallel to the 
long axis of the endocarp. 

Specimens.—OMSI Pb565, 586, 764, 1108, 1109, 
1128, 1322, 1323, 1502, 1636, UF 9285. (Lectotype: 
USNM 387567 from Florissant, Colorado [Lesquer- 
eux, 1883, pl. 32, fig. 21]). 

Discussion.— Cedrelospermum lineatum samaras are 
Preserved in the Nut Beds as impressions in siltstone. 
They were recovered only from a 1 meter-wide area 
Of the leaf-bearing horizon at the base of Face 3 in a 
Concentration of winged fruits that also included De- 
viacer. Cedrelospermum is known from nearly all Clar- 
Do paleobotanical localities. For comparative purpos- 
€s, the dimensions in the above description are 
deliberately limited to specimens from the Nut Beds. 
In the large size of endocarp and overall fruit length, 
this material conforms to C. lineatum, a species based 
Upon specimens from Florissant, Colorado (Manches- 
ter, 19892). Fruits from other localities of the Clarno 
Formation (e.g., West Branch Creek, Gosner Road, 
Teater Road) are smaller and appear to belong to an- 
Other species, C. nervosum, which is based upon ma- 
terial from the Green River Formation of Colorado, 
Utah and Wyoming (Manchester, 1989a). 

Cedrelospermum Saporta is an extinct genus of UI- 
Maceae known from the late Eocene to late Miocene 
Of Europe and the middle Eocene to Oligocene of west- 
ern North America (Manchester, 1987b, 1989a, c). Re- 
Cently, the genus has also been confirmed from Neo- 
&ne strata near Tepexi de Rodriguez, Puebla, Mexico 
(Magallón-Puebla and Cevallos-Ferriz, 1992), indicat- 
Mg a southern refugial population. Cedrelospermum 
Nervosum and C. lineatum fruits have been found at- 
tached to fossil twigs bearing narrow Zelkova-like leaves 
Ш the Eocene of Utah and Colorado (Manchester, 
19893), Characters of the fruits and foliage and of at- 
tached staminate flowers and pollen establish clear af- 
finities with the Ulmoideae (Manchester, 1989a, c). 


Family VITACEAE Jussieu 


The Vitaceae (grape family) include eleven extant 
Senera and about 800 species of lianas distributed in 
tropical to warm temperate regions. Seeds of this fam- 
Uy are well represented in the Tertiary of the northern 

*misphere as reviewed by Kirchheimer (1938), Tiff- 
Ney and Barghoorn (1976) and Cevallos-Ferriz and 
Stockey (1990). 

The Vitaceae are diverse in the Nut Beds, repre- 

Senting at least four genera: Ampelocissus (2 spp.), Am- 


pelopsis, Parthenocissus (2 spp.) and Vitis (2 spp.).* For 
the descriptions that follow, I use the terminology rec- 
ommended by Tiffney (1976), with the exception that 
the terms chalaza-base groove and chalaza-apex groove 
are reversed so that base refers to the hilar/micropylar 
end of the seed and apex to the opposite, rounded end. 
Extant genera overlap to some extent in seed mor- 
phology, and the evaluation of fossil remains is ham- 
pered by the lack of a thorough systematic work on 
seed morphology indicating characters that are diag- 
nostic at the generic level. Some morphological pat- 
terns appear to be confined to a single genus, while 
others occur in more than one modern genus. 


Genus AMPELOCISSUS Planchon 


Ampelocissus auriforma sp. nov. 
Plate 44, figures 16-18, 22 


Parthenocissus? sp., Scott, 1954, p. 82 , pl. 16, fig. 15. 


Etymology.—Aurus (L = ear) + forma (L = shape) 
referring to the shape of the enlarged ventral infolds. 

Description.—Seed cordiform in face view, epsilon- 
shaped in cross section, bilaterally symmetrical, base 
pointed, apex emarginate. Internal mold of seed coat 
length 3.5-7.1, avg. 5.4 mm (SD=1.04, n=28), width 
3.2—6.2, avg. 4.9 mm (SD=0.93, n=28), thickness 2.0— 
3.8, avg. 2.9 mm (50-0.49, n=26); dorsal surface con- 
vex, with a central elliptical to circular depression rep- 
resenting the chalaza, and a pronounced median groove 
passing from the base to the apex and over to the apex 
of the ventral surface; faint ruminations radiating from 
the chalaza; ventral infolds represented by a pair of 
large, smooth, elliptical, cup-like cavities separated by 
a thin median longitudinal raphal septum: lateral limbs 
of the seed 0.5-0.7 mm thick; seed coat ca 125 um 
thick, with a uniseriate layer of anticlinally oriented 
columnar cells. 

Specimens.—Holotype: UF 6572. Paratypes: HU 
59960, UCMP 10568-10578, UF 6543, 6571, 9524, 
9620-9624, UM 29935, 39564 (5 specimens), USNM 
355082, 355083 (10 specimens), 434987434997. 

Discussion.— This species is relatively common in 
the Nut Beds. Specimens are usually preserved as silica 
molds of the inside of the seed coat (Pl. 44, figs. 16, 
17. These internal seed coat molds are commonly re- 
covered within sedimentary molds of the outer seed 
surface. Such molds of the ventral seed surface have a 
distinctive bilobed appearence due to infilling of the 
pair of ventral infolds (Pl. 44, fig. 18). Scott (1954) 
figured and described specimens of this species as Par- 
thenocissus? sp. and interpreted the specimens to rep- 
resent detached dorsal sides of seeds similar to P. an- 
gustisulcata. However, transverse sections show that 


* For new information on wood, see note added in proof, p. 200. 


94 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


the seed coat is intact on both the dorsal and ventral 
sides and that the concave aspect ofthe ventral surface 
is an original feature (Pl. 44, fig. 22). 

The large cup-like ventral infolds of Ampelocissus 
auriforma distinguish it from all other Nut Beds Vi- 
taceae and are useful in generic placement. Seeds with 
comparably large infolds have been observed in some 
of the extant species of Ampelocissus, for example: A. 
martinii Planch. from the Philippines, A. imperialis 
from Java and A. costaricensis Lundell from Costa 
Rica. Although reports of Ampelocissus from the North 
American fossil record are rare, the genus was probably 
widely distributed in the Eocene; a single specimen 
preserved as a bilobed mold similar to this species is 
known from the Middle Eocene Green River Forma- 
tion of Utah (UF 8207, loc. 15755). Cevallos-Ferriz 
and Stockey (1990) reported another species based upon 
anatomically preserved seeds from the Middle Eocene 
Princeton chert of British Columbia. The Princeton 
chert species lacks the enlarged ventral infolds. Am- 
pelocissus is a genus with about 95 extant species, of 
tropical distribution. 


Ampelocissus scottii sp. nov. 
Plate 44, figures 11-15 


Etymology.—This species is named after Richard A. 
Scott. 

Description. —Seed cordate, bilaterally symmetrical; 
length 5.9, 5.9, 6.0 mm, width 4.8, 5.4, 6.0 mm, thick- 
ness 2.7, 2.7, 2.8 mm; ventral infolds short, about Уз 
as long as the seed, nearly parallel, slightly divergent 
apically, becoming wider in the apical Уз of the length; 
chalaza circular, central; chalaza-apex groove broad, 
chalaza-base groove indistinct, dorsal surface convex, 
with a central elliptical to circular chalaza. Internal 
mold of seed coat length 4.9-5.1 mm, width 3.9-5.4 
mm, thickness 2.2—3.5 mm; dorsal surface convex, with 
a central elliptical to circular depression representing 
the chalaza, and a pronounced median groove passing 
from the base to the apex and over to the apex of the 
ventral surface; faint ruminations radiating from the 
chalaza. 

Specimens.— Holotype: USNM 424665, seed. Para- 
types: UF 6545, USNM 355083, 355119, 434984, 
435118, seeds; UF 6542, 6544, internal molds of seed 
coat. 

Discussion.—Ampelocissus scottii resembles A. au- 
riforma in shape, size and morphology of the apex, 
base and dorsal surface. However, instead of the large 
cup-like ventral infolds, this species has short narrow 
divergent infolds (Pl. 44, fig. 11). Lacking the distinc- 
tive character of large ventral infolds, which to my 
knowledge is confined to Amplelocissus, the assign- 
ment of this species to the genus is less secure. How- 
ever, not all extant species of Ampelocissus have en- 


larged infolds. With its rugulate margins, broad chalaza- 
apex groove, and short divergent infolds, this fossil 
resembles the seeds of some extant species of Ampe- 
locissus, such as A. arachnoideus (Haask.) Planch. 

In general form, specimens of this species resemble 
those of Ampelopsis rooseae (Pl. 44, figs. 6-10), but 
they are much larger, more compressed dorsiventrally, 
and do not possess a chalaza-apex ridge. 


Genus AMPELOPSIS Michaux 


Ampelopsis rooseae sp. nov. 
Plate 44, figures 6-10, 21 


Tetrastigma, Bones, 1979, pl. 4, fig. 10. 


Etymology.—This species is named for Carrie L. 
Roose, acknowledging her technical assistance in this 
investigation. 

Description.—Seed subglobose, bilaterally symmet- - 
rical, rounded-triangular in dorsal and ventral views 
with smooth contours, slightly pointed at the base (hi- 
lum) in complete seeds, apex emarginate to broadly 
rounded, length of seed casts with intact seed coat 3.0- 
4.3, avg. 3.6 mm (SD=0.36, n=8), width 2.8-3.7, avg 
3.1 mm (SD=0.31, n=8), thickness 2.0-2.7, avg. 2.9 
mm (SD-0.23, n=8); chalaza typically spatulate, oC- 
casionally round, located in the central one-half to api- 
cal one-third of the seed on the dorsal surface; dorsal 
surface smooth or slightly ruminate; chalaza-bast 
groove present, narrow; chalaza-apex groove absent on 
the seed surface, although sometimes present on in- 
ternal molds of the seed coat; chalaza and apex typi- 
cally joined by a ridge that passes over the apex and 
joins with the raphe ridge; ventral infolds long and 
straight, diverging apically, becoming wider in upper 
third of their length, extending from a position well 
above the seed base and terminating approximately % 
of the length of seed toward the apex; groove corre- 
sponding to raphe preserved on the raphe ridge in com- 
plete seed casts; seed coat 130-150 um thick. Internal 
mold of seed coat sharply pointed basally, length 2) s 
4.5, avg. 3.3 mm (SD=0.41, n=36), width 2.3-4.1, av£: 
3.0 mm (SD=0.37, n=36), thickness 1.5-2.8, avg. 2 
mm (SD-0.30, n=36). 

Specimens.— Holotype: UF 6536, seed cast. Рага- 
types: UCMP 10596-10597, UF 6537, 6538, 9575, 
9659, 9660, 9661, USNM 312768, 312769 (Bones, 
1979, pl. 4, fig. 10), 355122 (2 specimens), seed casts: 
UCMP 10598-10606, UF 6538, 9662-9691, USNM 
355090 (20 specimens), 355093 (106 specimens), in- 
ternal molds of seed coat. 

Discussion.— This species is similar in size to the 
seeds of Vitis tiffneyi but it is distinguished by its Ven” 
tral infolds, which are longer and divergent, rather tha? 
parallel, and by the well-developed raphe ridge that 
typically passes over the apex to join with the chalaza- 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 95 


Most specimens are globular, suggesting one-seed per 
berry, but in some of the seeds the ventral surface has 
flattened facets suggesting more than one seed per ber- 
гу. 
Seeds with short divergent ventral infolds occur in 
Some of Ше extant species of Ampelopsis and Vitis. 
Whereas Vitis seeds generally have a chalaza-apex 
groove, those of Ampelopsis lack such a groove and 
typically have a ridge in the same position, as does this 
fossil. A. rooseae bears a close resemblance to Ampe- 
lopsis monasteriensis Kirchheimer (= A. rotundata Reid 
et Chandler 1933 auct non Chandler 1926; Chandler 
1961b) from the Eocene London Clay flora. A com- 
Pression specimen that appears to be identical to this 
Nut Beds species is known from the Eocene Green 
ү Formation near Watson, Utah (UF 8206, loc. 
5755). 


Genus PARTHENOCISSUS Planchon 


Parthenocissus angustisulcata Scott 
Plate 45, figures 6, 7 


Р, arthenocissus angustisulcata Scott 1954, p. 81, pl. 16, fig. 14. 
Vitaceae. Undetermined genus, Bones, 1979, pl. 4, figs. 11, 12. 


Description. —Seed cast subglobose, obovate in face 
View, bilaterally symmetrical to asymmetrical, length 
5.4-6.4, avg. 6.0 mm (SD=0.45, n=3), width 4.6-5.5, 
àvg. 5.1 mm (SD=0.45, n=3), thickness 2.9-4.3, avg. 
3.5 mm (SD=0.74, n=3), surface usually smooth, but 
Undulating in some specimens, ventral infolds long, 
Slightly curved, diverging apically, extending from the 
Very base of the seed to the apical % of the seed, raphe 
Паре triangular, slightly raised above the seed body, 
Chalaza circular to spatulate, centrally positioned, faint 
Tuminations radiating from chalaza over the dorsal 
Surface; chalaza-base groove and chalaza-apex groove 
Well developed. 

Specimens.— Holotype: UM 29934 (Scott, 1954, pl. 
16, fig. 14). Hypotypes: UF 6544, 9737, 9738, 9880- 
9882, USNM 355128, 434986. 

Discussion.— This species was described by Scott 
(1954) on the basis of a single specimen showing the 
Ventral side only. More complete seeds now available 
also show the dorsal surface, with a pronounced cha- 
aza-base groove (Pl. 45, fig. 7) and give a better un- 

*rstanding of the variability within the species, en- 
abling the more complete description provided above. 

5 Scott noted, important features of Parthenocissus 
angustisulcata include the deep apical notch marking 

е course of the raphe, and long, narrow, divergent 
Ventral infolds (e.g., Pl. 45, fig. 6). 

he long narrow, apically divergent ventral infolds 
And narrow raphe ridge were used to place this in Par- 
Menocissus. Scott noted that it differs from the modern 
Species and from P. monasteriensis (Reid et Chandler) 


Scott ofthe London Clay flora by its larger size. Wheth- 
er this size difference and the pronounced apical cleft 
are admissable within Parthenocissus is open to further 
investigation. 


Parthenocissus clarnensis sp. nov. 
Plate 45, figures 1—5 


Etymology.— The epithet refers to the Clarno For- 
mation. 

Description.—Seed subovoid, obovate in face view; 
cross section rounded in some, rounded-triangular in 
others, due to variable development of ventral facets, 
bilaterally symmetrical to asymmetrical; length 3.0- 
4.8, avg. 3.8 mm (SD=0.45,n=25), width 2.8-4.2, avg. 
3.4 mm (SD=0.36, n=25), thickness 2.0-3.5, avg. 2.2 
mm (SD-0.34, n=25); ventral infolds long, slightly 
curved, diverging apically, extending from the very 
base of the seed to the apical У of the seed, raphe ridge 
triangular slightly raised above the seed body, surface 
usually smooth, but undulating in some specimens, 
chalaza typically spatulate, occasionally round, posi- 
tioned central or somewhat offset toward the apex, 
radiating undulations usually week, chalaza-base groove 
absent or faint, chalaza apex-groove present, weak 
(stronger in the globose specimens, weaker in the fac- 
eted ones); seed coat 200—300 um thick, with a uni- 
seriate layer of columnar cells 200 um high and 12-16 
um thick and an outer layer 40 to 80 um thick com- 
posed of isodiametric cells ca. 14-18 um in diameter. 

Specimens.— Holotype: UF 6539. Paratypes: UCMP 
10584, 10595, UF 6540, 6541, 9583, 9693-9717, 
USNM 355085 (19 specimens), 355087 (7 specimens), 
355088 (12 specimens), 355089 (12 specimens), 355091 
(12 specimens) 355100 (28 specimens), 355134 (8 
specimens). 

Discussion.— Parthenocissus clarnensis is character- 
ized by long, divergent ventral infolds, a lack of a cha- 
laza-base groove and weak to well-developed chalaza- 
apex groove (Pl. 45, figs. 1-4). Some of the specimens 
are relatively globose and probably were borne one per 
fruit. Others have one or two flattened faces on the 
ventral side suggesting that they were borne two or 
more per fruit. It is much more common than P. an- 
gustisulcata, which has larger seeds with more pro- 
nounced chalaza-base and chalaza-apex grooves. 

This species resembles Parthenocissus monasterien- 
sis (Reid et Chandler) Scott from the London Clay flora 
and is similar to extant P. vatascea from northern Mex- 
ico and Vitis amurensis J.Y. Li from Siberia. 


Genus VITIS L. 


Vitis magnisperma Chandler 
Plate 45, figures 8-13 


Vitis magnisperma Chandler 1961b, p. 247, pl. 24, figs. 29, 30. 
Vitis, Bones, 1979, pl. 4, fig. 9. 


96 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Description. — Internal mold of seed coat more ог less 
elliptical in face view, truncate or emarginate apically 
with a slight to well-developed chalaza-apex groove, 
chalaza-base groove faint or absent; length 7.0-10.3, 
avg. 8.9 mm (SD=1.09, n=6), width 5.5-6.9, avg. 6.2 
mm (SD=0.57, n=6), thickness 2.8-3.0, avg. 2.9 mm 
(SD=0.10, n=4); ventral infolds long, somewhat di- 
vergent apically, straight to curved, extending from the 
extreme base to the apical Y, of seed; chalaza circular 
to cordate, centrally positioned; faint ruminations ra- 
diate around the chalaza and over the dorsal surface; 
chalaza-apex groove well developed, chalaza-base 
groove faint or absent. 

Specimens. — UCMP 10607, UF 9735, 9879, USNM 
312761 (Bones, 1979, pl. 4, fig. 9, right), 355095, 
355098, 355127 (Bones 1979, pl. 4, fig. 9, left), 355130, 
434985. 

Discussion.— This species includes the largest vita- 
ceous seeds known from the Nut Beds, some a full 
centimeter in length. Only internal molds of the seed 
coat are represented, so the full size of the seed was 
somewhat larger. With its elongate ventral infolds, 
shallow chalaza-apex groove and ruminations ema- 
nating from the chalaza, this species does not differ in 
any essential characters from those in the London Clay 
described as Vitis magnisperma Chandler (Chandler, 
1961b). These seeds are larger than those typical of 
extant Vitis species, and the species might represent 
another genus. However Chandler's assignment is re- 
tained pending further comparative work with extant 
Vitaceae. 


Vitis tiffneyi sp. nov. 
Plate 44, figures 1—5, 19, 20 


Etymology.— This species is named for Bruce H. 
Tiffney in recognition of his contributions to paleo- 
botany and the study of fossil vitaceous seeds. 

Description.—Seed subglobose, bilaterally symmet- 
rical, with smooth contours, pointed at the base (hi- 
Ium), apex slightly emarginate; length of complete seed, 
with seed coat intact 3.7—4.0, avg. 3.9 mm (SD=0.14, 
n=5), width 3.2—3.5, avg. 3.4 mm (SD=0.13, n=5), 
thickness 2.3-2.7, avg. 2.5 mm (SD=0.14, n=5); cha- 
laza circular to elliptical, positioned centrally on the 
dorsal surface, sometimes surrounded by weak radi- 
ating undulations; chalaza-base groove moderate, cha- 
laza-apex groove pronounced, extending over the seed 
apex and merging with the raphal ridge of the ventral 
side; paired ventral infolds straight, short, approxi- 
mately half the length of the seed, oriented nearly par- 
allel to one another; with a fine groove at surface of 
raphe ridge; raphe ridge widening only slightly, if at 
all, toward the apex; seed coat 90-100 um thick, with 
a uniseriate layer of columnar cells ca. 90 um high and 
15-20 um wide. Internal mold of seed coat morpho- 


logically similar to the seed surface but with sharp base, 
emarginate apex, length 2.8-3.8, avg. 3.3 mm 
(SD=0.24, n=40), width 2.4-3.3, avg. 2.9 mm 
(SD=0.19, n=40), thickness 1.3-10 2.8, аур. 2.0 mm 
(SD=0.27, n=40). 

Specimens.—Holotype: UF 6534, seed cast. Para- 
types: UF 9573, 9625-9628, complete seed casts; HU 
60070 (Tiffney and Barghoorn, 1976, pl. II, J), UCMP 
10579-10583, UF 6563, 6535, 9572, 9629-9658, 
USNM 355094 (26 specimens), 355102 (20 speci- 
mens), internal molds of seed coat. 

Discussion. — The distinctive features of this seed type 
are the parallel, relatively short ventral infolds, typi- 
cally circular chalaza, bisymmetry and pronounced 
chalaza-apex groove. The species is abundant in the 
Nut Beds, represented both by complete seed casts with 
the seed coat intact (Pl. 44, fig. 1, 2) and by molds of 
the inside of the seed coat (Pl. 44, figs. 4, 5). Some of 
the complete seed casts have a portion of the seed coat 
fractured away, showing a relatively thin seed coat, 
and revealing the internal mold of the seed coat (Pl. 
44, fig. 3). The internal molds, most abundant at the 
locality, are distinguished by the more sharply pointed 
hilar projection and by the outline of broken testa suf- 
rounding each of the ventral infolds. In general size 
and outline, these specimens can be confused with those 
of Ampelopsis rooseae, however, the latter are distin- 
guished by having divergent (Pl. 44, figs. 6, 8, 10), 
rather than parallel (Pl. 44, figs. 1, 4) ventral infolds 
and a thicker seed coat (cf. Pl. 44, figs. 3, 7, 19, 21). 

This species is similar in shape and size to V. subgl- 
obosa Reid et Chandler from the London Clay (Reid 
and Chandler, 1933; Chandler, 1961b). Tiffney and 
Barghoorn (1976) examined this Clarno species and 
noted that the short, wide ventral infolds indicate sim- 
ilarity to seeds of extant Ampelopsis and to many Pa- 
leogene species, including Vitis subglobosa Reid et 
Chandler, Ampelopsis crenulata, A. monasteriensis (A. 
rotundata emend. Chandler 1960), Vitis platyformis. 
V. rectisulcata, Palaeovitis paradoxa Reid et Chandler, 
V. obovoidea Chandler. Affinities to Ampelopsis prob- 
ably can be ruled out by the absence of a well-devel- 
oped chalaza-apex ridge. Based on their examination 
of this species from the Nut Beds, Tiffney and Bar- 
ghoorn (1976, p. 185) observed that the “The com- 
bination of distinctive infolds with other Vitis char- 
acters leads one to suspect that they represent either 
an unrecognized modern form or an extinct lineage 10 
the genus Vitis." 


INCERTAE SEDAE 
Genus ANKISTROSPERMA gen. nov. 
Etymology.— Ankistron (Gr = fish hook) + sperm 
(Gr — seed), referring to the J-shape of the seed. 
Type species. —Ankistrosperma spitzerae sp. nov. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 97 


Ankistrosperma spitzerae sp. nov. 
Plate 46, figures 11-13 


Etymology.—The epithet recognizes contributions 
of Josephine Spitzer to this project. 

Description.—Cast J-shaped, often bilaterally sym- 
metrical, circular in cross section, pointed at short end, 
blunt or truncate at long end; length 2.0-2.9 mm, width 
across plane of symmetry 0.7-0.9 mm, constricted in 
the apical region into a cylindrical neck 0.3-0.5 mm 
long, surface smooth. 

Specimens.—Holotype: UF 9806. Paratypes: UF 
9807-9814, 9815 (10 specimens), USNM 355568 (22 
Specimens), 355571. 

Discussion.—This species is based upon distinctive 
J-shaped casts that appear to represent seeds with a 
curved embryo and a constricted, neck-like hilar area 
(Pl. 46, figs. 11-13). The short, pointed end might rep- 
resent the micropyle. Affinities with respect to modern 
families are unknown. 


Genus ANONYMOCARPA gen. nov. 


Etymology.—Anonymos (Gr = nameless, unknown) 
+ karpos (Gr = fruit), referring to the uncertain familial 
affinities of this fruit. 

Type species. —Anonymocarpa ovoidea sp. nov. 


Anonymocarpa ovoidea sp. nov. 
Plate 45, figures 14-17 


Etymology.— Ovatus (L = egg-shaped) + oid (Gr = 
like, resembling), referring to the endocarp shape. 
| Description. — Fruit ellipsoidal, circular in cross sec- 
tion, rounded apically and basally, unilocular, single- 
Seeded; length 29 mm, equatorial diameter 22 mm, 
репсагр consisting of three layers: 1) inner layer 120- 
140 um thick, uniseriate, composed of anticlinally 
elongate cells 120-140 um high, 20-30 um wide, 2) 
Middle layer 400 um thick composed of longitudinally 
Oriented fibers, 50-100 um wide, 3) outer layer about 
400 um thick composed of narrow, horizontal, peri- 
Clinally oriented fibers, this outer layer with prominent 
Qngitudinal vascular bundles up to 200 um in diam- 
eter, 

Specimen.— Holotype: USNM 355614. 

Discussion.—This species is known from one fruit 
that has been sectioned transversely to show the single 
locule and wall structure. Although the fruit is ana- 
tomically distinctive, its systematic affinities remain 
Obscure. 


Genus ASCOSPHAERA gen. nov. 


Etymology.—Asco (Gr = bag, bladder) + sphaero 
(Gr = ball), referring to the sack-like form of the fruit 
апа spheroidal form of the seed. 

Туре species.— Ascosphaera eocenis sp. nov. 


Ascosphaera eocenis sp. nov. 
Plate 46, figures 6-10 


Etymology.— The epithet refers to the Eocene age of 
the Nut Beds deposit where this species occurs. 

Description.— Fruit bulb-shaped, radially symmet- 
rical, unilocular, single-seeded, circular in cross sec- 
tion, length 3.1-4.4 mm, width 2.0-3.0 mm, surface 
rough; fruit wall 0.4-0.6 mm thick with an outer layer 
of polygonal cells 100-200 um in diameter. Locule cast 
spheroidal, length 2.0-2.1 mm, equatorial diameter 
1.7-1.8 mm, smooth, with a circular truncation at the 
base, and a slight rise at the apex, sometimes with a 
faint longitudinal groove (raphe?). 

Specimens.— Holotype: UF 6318. Paratypes: UF 
6319, 6320, 9789, 9790, 9873, fruits; UF 6343, 6415, 
9770, USNM 424753, 424759, 424789, locule casts. 

Discussion.— This species is represented by silicified 
fruits shaped like light bulbs (Pl. 46, figs. 6, 7) and 
globose locule casts (Pl. 46, figs. 8, 9). A fractured fruit 
with the locule cast partially exposed (Pl. 46, fig. 7) 
helps to establish the link between the complete fruits 
and isolated locule cast specimens. 


Genus AXINOSPERMA gen. nov. 


Etymology.—Axine (Gr = Ax, wedge) + spermum 
(Gr = seed). 
Type species. — Axinospermum agnostum sp. nov. 


Axinosperma agnostum sp. nov. 
Plate 46, figures 1—5. 


Etymology.—Agnostos (Gr = unknown, unknow- 
ing), alluding to the unknown affinities of this taxon. 

Description.—Seed bilaterally symmetrical, more or 
less elliptical with rounded base and apex as seen in 
lateral view; laterally compressed with a median keel 
developed apically and ventrally in the plane of sym- 
metry, length 1.9-2.2 mm, width in the plane of sym- 
metry 1.2-1.6 mm, thickness across the plane of sym- 
metry 0.7-1.0 mm, surface smooth, hilum subapical 
and ventral in the plane of symmetry; embryo cavity 
elongate, curved ventrally, circular in cross section. 

Specimens.— Holotype: UF 6290. Paratypes: UF 
9799, 9800, 9816-9829, USNM 424711-424713, 
424716, 424736. 

Discussion.— These small seeds are preserved as sil- 
ica casts. The laterally flattened shape and the keel 
along the apical and dorsal margins in the plane of 
symmetry are distinctive features. Sometimes the api- 
cal crest is broken away, revealing a lobe ofthe embryo 
cavity (Pl. 46, figs. 3, 4). A transversely sectioned seed 
(Pl. 46, fig. 5) reveals the elliptical cross section of the 
embryo cavity. 


98 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Genus BONESIA gen. nov. 


Etymology.—Named in memory of Thomas J. Bones, 
the collector. 
Type species.— Bonesia spatulata sp. nov. 


Bonesia spatulata sp. nov. 
Plate 47, figures 1-10 


Etymology.— Spatula (L = diminutive of spatha, a 
broad blade, paddle for stirring) referring to the spat- 
ulate shape of the locule casts. 

Description. — Fruit unilocular, single-seeded, wide- 
elliptical in face view, dorsiventrally flattened, ellip- 
tical or lenticular in cross section; quadrilaterally sym- 
metrical about the longitudinal axis, base and lateral 
margins rounded, apex mucronate, surface smooth with 
a low-rounded, median longitudinal ridge extending 
from the base to the apex on both dorsal and ventral 
sides; pericarp 0.5 mm thick, composed of an inner 
layer of radial files of isodiametric sclereids 50-63 um 
in diameter and a uniseriate outer layer of anticlinally 
oriented columnar sclerenchymatous cells 83 um high, 
12-15 um wide. Locule cast smooth, similar in shape 
to endocarp, with a circular basal depression approx- 
imately 1 mm in diameter, locule cast length 4.3-6.2, 
avg. 5.0 mm (SD=0.5, n=14), width 3.9-5.2, avg. 4.5 
mm (SD=0.39, n=14), thickness 1.4-2.4, avg. 1.9 mm 
(SD=0.31, n=14); seed with a thin shriveled seed coat 
and large embryo with a pair of planar cotyledons. 

Specimens.— Holotype: USNM 424642. Paratypes: 
UCMP 10728, 10729, UF 6355, 8583, 8584, 9398- 
9404, 9472, USNM 354057 (3 specimens), 424643, 
435088. 

Discussion.— Bonesia spatulata fruits usually are 
preserved as white translucent chalcedony locule casts 
that are readily identified by their dorsiventral flatten- 
ing, slightly raised median dorsal and ventral ridges, 
slight apical beak (Pl. 47, figs. 1-4) and prominent basal 
circular scar (Pl. 47, figs. 4—6). Occasionally part of the 
fruit wall is preserved and may be sectioned to reveal 
a relatively thick endocarp with an outer uniseriate 
columnar layer (Pl. 47, figs. 8, 10). Sections through 
the locule casts reveal a single seed within the locule, 
with a pair of large, planar cotyledons (Pl. 47, figs. 8, 
9). 


Genus COMMINICARPA gen. nov. 


Etymology.— Comminus (L = near, at close quar- 
ters) + karpos (Gr = fruit), referring to the close spacing 
of fruits within the infructescence. 

Type species. — Comminicarpa friisae sp. nov. 


Comminicarpa friisae sp. nov. 
Plate 48, figures 1—7 


Etymology.— The epithet recognizes the contribu- 
tions of Else Marie Friis to paleobotany. 


Description.—Infructescence spike-like, at least 16 
mm long, and 10 mm in diameter consisting of nu- 
merous achenes closely packed together on an elongate 
axis, perianth not evident. Fruits apparently grouped 
in threes, more or less obovate, laterally faceted and 
angular-subrounded in cross section through crowding 
effect of neighboring fruits, cuneate in longitudinal sec- 
tion, narrowed basally, apex truncate-rounded with 
central protruding style; fruit length 2.5-2.8 mm, width 
0.85-1.1 mm; pericarp 80-100 um thick, composed 
mostly of isodiametric cells about 10-15 um in di- 
ameter, but with an intermittent middle uniseriate lay- 
er of large rectangular to subrounded cells 30-50 ит 
in diameter that appear to be idioblasts. Fruits uni- 
locular with a single seed pendulous from the apex. 
Seed coat consisting of a thin layer of narrow (5 um) 
axially elongate cells. 


Specimens.— Holotype: USNM 355697. Paratypes: . 


UF 9349, 9582, 9874-9878. 

Discussion.— This species is represented by one in- 
fructescence (Pl. 48, figs. 1—3, 5) and several specimens 
with isolated achenes in close proximity scattered in 
small pieces of sediment. The fruits are achenes, each 
with a prominent central style. The seed coat is well 
preserved and appears to have been well cutinized. 


Genus CUNEISEMEN gen. nov. 


Etymology.— Cuneus (L = wedge) + semen (L = 
seed). 
Type species. — Cuneisemen truncatum sp. nov. 


Cuneisemen truncatum sp. nov. 
Plate 47, figures 11-16 


Etymology.— Truncus (L — cut off), referring to the 
apical facet. 

Description. —Seed/pyrene elongate, typically bisy- 
metrical, shaped like a lemon segment, with a rounded 
dorsal face and a pair of relatively flat ventral faces 
meeting at an angle of ca. 50—60°, giving a rounded- 
triangular cross section, truncate apically with a tri- 
angular facet, ca. 0.9-1.2 mm in diameter, bluntly 
pointed at the base; length 4.0-4.8, avg. 4.6 mm 
(SD=0.60, n=9), width 2.2-3.1, avg. 2.5 mm (SD=0.30, 
n=9), dorsiventral thickness 2.1-2.5, avg. 2.3 mm 
(SD=0.13, n=9), dorsal and ventral faces very finely 
striate longitudinally; outer coat thin, terminating api- 
cally in a rim around the truncation. 

Specimens.— Holotype: UF 6392. Paratypes: UCMP 
10687, UF 9362, 9364-9368, 9724. 

Discussion.—This species is represented by wedge- 
shaped chalcedony casts that appear to represent pY- 
renes or seeds. The angular shape suggests that they 


may have been borne close together in the fruit. of 


eight specimens, seven are bilaterally symmetrical, with 
the plane of symmetry bisecting the ventral angle (Pl. 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 99 


47, figs. 11, 15) and one is asymmetrical (Pl. 47, fig. 
16), possibly distorted through ontogeny. The outer 
Covering (seed coat or exocarp?) forms a rim around, 
but does not cover, the apical truncation (Pl. 47, fig. 
14), indicating that the truncation is probably the at- 
tachment scar. The familial affinities of this taxon re- 
Main undetermined. 


Genus DENTISEMEN gen. nov. 


Etymology.— Dentis (L = tooth) + semen (L = seed) 
Type species.— Dentisemen parvum sp. nov. 


Dentisemen parvum sp. nov. 
Plate 46, figures 14—17 


Etymology.—Parvus (L = little), referring to the small 
Seed size. 

Description.—Seed cast trigonal in outline with flat 
base and pointed apex, rounded dorsally, bilaterally 
Symmetrical, with a median longitudinal infold or slit 
On the ventral surface in the plane of symmetry ex- 
tending from the apex to a point about !^ of the distance 
from the base; length 2.5-2.7 mm, width 1.4-1.7, 
thickness 1.0-1.3 mm; surface micro-verrucate. 

Specimens.—Holotype: UF 9766. Paratypes: UCMP 
10738, UF 9767, USNM 355434, 355670. 

Discussion.—These appear to be internal molds of 
Seed coat. They are preserved in white chalcedony, 
Without cellular detail. Although distinctive in the flat 
base, median ventral slit (Pl. 46, figs. 14, 16) and ver- 
Tucate surface (Pl. 46, figs. 16, 17), the affinities among 
extant families remain unknown. 


Genus DUROCARPUS gen. nov. 


Etymology.— Durus (Г. = hard, tough, lasting) + kar- 
Pos (Gr = fruit). 
Type species.— Durocarpus cordatus sp. nov. 


Durocarpus cordatus sp. nov. 
Plate 48, figures 8-13 


Etymology.—The epithet, cordatus (L = heart- 
Shaped) refers to the seed outline. 
. Description. —Fruit wide obovate in face view, el- 
liptical in cross section, quadrilaterally symmetrical 
with the two planes of symmetry intersecting at right 
angles in the vertical axis; in face view apex more or 
ss flat, base wide-cuneiform; length 5.5 mm, width 
6.0 mm, thickness 3.9 mm; bicarpellate and bilocular 
With one seed per locule; surface smooth and rounded; 
Pericarp thick, 1.2 mm, septum 2 mm thick; locules 
Obovate, 1.8 mm high, 0.8 mm wide, placentation ven- 
tral near the apex of locule, ovules pendulous; a central 
Column of vascular strands passing medially from the 
base to the apex, tapering toward the apex; locule lining 
апа endocarp composed of a four- to ten-seriate layer 
9f thick-walled isodiametric cells 25-60 um in diam- 


eter; mesocarp, forming the bulk oftissue between and 
surrounding the two endocarps, composed of thin- 
walled cells 42-75 um in diameter, with islands of 
thick-walled cells similar to those of the endocarp, 
prominent branching vascular strands and lacunae filled 
with dark contents, exocarp not preserved. 

Specimen.—Holotype: UF 6110. 

Discussion. — Durocarpus is represented by a single 
specimen that is a distinctive bilocular, relatively thick- 
walled fruit. The specimen was sectioned longitudi- 
nally (Pl. 48, fig. 10), revealing the ventral, subapical 
placentation (Pl. 48, figs. 11, 12), then reassembled and 
sectioned transversely (Pl. 48, fig. 13). The fruit anat- 
omy is well preserved. Extant families with similar 
characteristics include Anacardiaceae, Euphorbiaceae, 
Meliaceae. No closely similar extant genera were ob- 
served among these families, however. 


Genus FERRIGNOCARPUS gen. nov. 


Etymology.— This genus is named for James P. Fer- 
rigno, recognizing his services to paleobotany at the 
U.S. National Museum. 

Type species.— Ferrignocarpus bivalvis sp. nov. 


Ferrignocarpus bivalvis sp. nov. 
Plate 49, figures 1-15, Text-figure 18 


Etymology.— Ві (L = two) + valva (L = leaf of a 
folding door), referring to the two-valved structure of 
the епдосагр. 

Description. — Fruit unilocular, endocarp elliptical in 
face view, lensoidal in cross section, quadrilaterally 
symmetrical with two planes of symmetry intersecting 
at right angles in the vertical axis, base and apex round- 
ed, endocarp length 4.5-9.0, avg. 6.3 mm (SD- 1.48, 
n=19), width 3.4—6.4, avg. 4.7 mm (SD=0.81, n=18), 
thickness 2.1-3.9, avg. 2.9 mm (SD=0.57, n=17), bi- 
valved, splitting along the major plane of symmetry, 
endocarp wall 1.0-1.2 mm thick, composed of: an in- 
ner layer about 50 um thick composed of square to 
rectangular cells up to 15 um in diameter, a middle 
layer typically 300 um thick of vertically oriented fibers 
25 um in diameter, and a thick outer layer of anticlin- 
ally oriented short fibers, 55-75 um long and 20-30 
um in diameter; rhomboidal crystals in idioblasts com- 
mon at the contact between the two layers of fibers. 
Locule cast smooth, elliptical to broadly obcordate in 
face view, lensoidal to elliptical in cross section, quad- 
rilaterally symmetrical, with a keel around the margin 
in the major plane of symmetry, and a shallow median 
longitudinal groove on both dorsal and ventral faces 
in the minor plane of symmetry; base slightly cordate 
or emarginate in face view, rounded in lateral view, 
sometimes with a thin straight stalk; apex obtuse in 
dorsiventral view, acute in lateral view. 

Specimens.— Holotype: USNM 424624. Paratypes: 


100 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


1ст 


Text-figure 18.— Ferrignocarpus bivalvis gen. еї 5р. поу. A, Ventral 
view of a single fruit valve showing thick wall and concave locule. 
B, Typical specimen in transverse section. 


UF 5669, 9386-9393, USNM 355348, 355369 (2 spec- 
imens), 355483 (8 specimens), 355404, 424803, 
424804. 

Discussion.— Ferrignocarpus bivalvis is represented 
mostly by locule casts that are distinctive in their cor- 
date outline with a longitudinal keel in the major plane 
of symmetry and a longitudinal median groove in Ше 
minor plane of symmetry (e.g., Pl. 49, figs. 2-4, 7). 
Only a few of the locule casts retain some or all of the 
thick endocarp wall (Pl. 49, figs. 5, 6, 8, 9). One spec- 
imen is a mold in sandstone representing one valve of 
the endocarp pressed into the sediment showing the 
cordate outline of the locule within the elliptical outline 
of the endocarp (PI. 49, fig. 1). The anatomy of the 
wall and locule contents are revealed in thin sections 
(Pl. 49, figs. 10-15). As seen in transverse sections, the 
locule contains an elliptical mass of parenchymatous 
tissue with a central 4-lobed space near the base (PI. 
49, figs. 12-14). The original structure is open to in- 
terpretation. The parenchymatous body may represent 
the storage tissue of a single seed with the 4-lobed 
central space representing the position of archegonia 
or of a decomposed embryo. The central part of the 
four-lobed space has an incomplete septum that ap- 
pears to give rise to two placentae (Pl. 49, fig. 14). 


. Genus FIMBRIALATA gen. nov. 
Etymology.— Fimbria (L = fringe) + alatus (L = 
winged). 
Type species.— Fimbrialata wingii sp. nov. 


Fimbrialata wingii sp. nov. 
Plate 50, figures 14-19; Text-figure 19 
Etymology.—This species is named after Scott L. 


Wing, recognizing his contributions to Eocene paleo- 
botany. 


3mm 


Text-figure 19.—Fimbrialata wingii gen. et sp. nov. A, Seed in 
lateral view. В, Seed in face view showing the encircling wing. С, 
Transverse section of seed. 


Description.—Seed subellipsoidal, asymmetrical, 
dorsiventrally compressed, consisting of a central seed 
body with a thin wing surrounding approximately in 
the dorsiventral plane; seed body more or less elliptical 
in cross-section, base and apex of seed and of seed 
body rounded in face view; seed height 2.9-3.7 mm, 
width 1.7-2.1 mm, thickness 0.6-0.9 mm; wing with 
striations oriented perpendicular to its margin, forming 
a flange about 0.3 mm wide around the seed body; 
surface of seed body smooth. Seed body with copious 
endosperm, embryo minute. 

Specimens.— Holotype: USNM 435057. Paratypes: 
UF 6286, 6289, 6867, USNM 435058. 

Discussion. — Because of the delicate nature of these 
fossils, the flanging wing often is partially broken. At 
first it appeared that the Fimbrialata wingii specimens 
might represent locule casts of a bivalved fruit with 
the “wings” representing the silica-filled crack along 
the dehiscence plane. However, transverse sections ге“ 
veal a cutinized layer outlining a well-defined wing (Pl. 
50, fig. 19). 


Genus FRAGARITES gen. nov. 


Etymology.— Fragaria (Г, = strawberry) + пез (Gr 
= like, having the nature of), referring to the similarity 
in shape to the achenes of Fragaria. 

Type species.— Fragarites ramificans sp. nov. 


Fragarites ramificans sp. nov. 
Plate 50, figures 1—4 


Etymology. — The epithet refers to the ramifying suf- 
face ribs. 


| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 101 


Description.—Seed (or achene?) bilaterally symmet- 
rical, laterally compressed and more or less D-shaped 
In the plane of symmetry, i.e., rounded in lateral out- 
line except for the straight hilar margin, and lenticular 
Ш cross section; length along hilar margin 2.4-3.7, avg. 
3.0 mm (SD=0.63, n=7), width 2.0-3.3, avg. 2.4 mm 
(SD=0.53, n=7), thickness 1.4-2.3, avg. 1.7 mm 
(SD=0, 39, n=7), length/thickness ratio 1.6-1.9; upper 
end of the flat hilar margin with a micropylar point, 
lower margin of the flat hilar margin rounded, surface 
Covered by a reticulum of bifurcating fibrous ribs ca. 
0.1 mm in diameter that are oriented approximately 
at right angles to the hilar edge. Inner mold of seed 
Coat similar in shape to the seed, with smooth surface. 

Specimens.— Holotype: USNM 424729. Paratypes: 
UCMP 10745, UF 6421, 9557-9564. 

Discussion.—Fragarites ramificans is represented by 
chalcedony casts of a seed or achene with prominent 
bifurcating ribs that run at right angles to the hilar edge 
(PL 50, figs. 1, 3). Most commonly, it is preserved as 
ап inner mold of the seed coat, with little or none of 
the outer reticulum adhering. Internal anatomy has not 
been determined. At least superficially, these speci- 
Mens with their D-shaped outline and transverse rib- 
bing resemble achenes of F ragaria in the Rosaceae. 


Genus GLOBULICARPIUM gen. nov. 
Etymology.— Globulus (L = bead) + karpos (Gr = 
ruit). 
Type species.— Globulicarpium levigatum sp. nov. 


Globulicarpium levigatum sp. nov. 
Plate 50, figures 5-13 


Etymology.— Laevis (L = smooth), referring to the 
Smooth surface of the fruit. 

Description. —Fruit nearly spherical, but with the 
Pedicel scar raised on a nipple-like projection, hence 
Slightly longer than wide, unilocular, single-seeded, in- 
dehiscent, length 4.3-6.8, avg. 5.5 mm (SD=0.79, 
113), equatorial diameter 3.2-5.7, avg. 4.5 mm 
(SD=0.66, n=17); surface smooth, sometimes shiny; 
Pericarp 0.9 mm thick, composed of at least four suc- 
Cessive layers: 1) locule lining composed of a uni- to 
biseriate layer about 24 um thick of thin-walled hor- 
‘Zontally oriented cells 10-15 um high by 50-87 um 
löng, 2) a thick layer, 400-500 um thick, consisting of 
ISodiametric to horizontally elongate thick-walled 
Sclereids varying to large size (50-210 um in diameter), 
Often with dark contents, nested among horizontally 
elongate parenchyma cells 15—40 um thick, 60—120 um 
Wide, this layer containing about 20 longitudinal vas- 
Cular bundles with helically thickened elements situ- 
ated about in the middle of the layer, the inner М of 
Ше middle layer in most specimens usually poorly pre- 
Served, detail obscured by the confluence of the opaque 


contents of oil cells or sclereids; 3) a band 150-175 
um thick of isodiametric parenchyma cells 15-25 um 
in diameter and, 4) a uniseriate outer layer of anti- 
clinally elongate cells 35 um high, 8-12 um wide. Loc- 
ule globose, smooth-walled, seed filling the locule, cot- 
yledons not preserved. 

Specimens.— Holotype: USNM 355663. Paratypes: 
UCMP 10623, UF 6349, 8804-8807, 9331, 9332, 
9506—9512, USNM 435009-435011, 435038. 

Discussion.— Globulicarpium levigatum is repre- 
sented in the Nut Beds by casts and permineralized 
fruits. Although similar in size and globose shape to 
Celtis which is also common in the Nut Beds, it is 
distinguished in the field by its smooth outer surface 
and nipple-like attachment scar. Transverse and lon- 
gitudinal sections have revealed the thickness and 
anatomy of the pericarp (Pl. 50, figs. 11-13). The peri- 
carp was probably leathery; a few specimens are shriv- 
eled like a raisin (e.g., UF 9332). 

The affinities of Globulicarpium could possibly be 
with the Lauraceae, but the lack of columnar cells, 
which are characteristic of the endocarps of Lauraceae 
(Reid and Chandler, 1933), and lack of preservation 
of large plano-convex cotyledons have made me hes- 
itant to assign it to that family. This species appears 
to be very similar in external morphology to a speci- 
men figured as “undetermined Lauraceae fruit" by 
Collinson (1983, fig. 216) from the London Clay flora, 
but I have not conducted a detailed comparison. 


Genus HEXACARPELLITES gen. nov. 


Etymology.— Hexa (Gr = six) + karpos (Gr = fruit), 
referring to the six carpels evident in this fruit. 
Type species. — Hexacarpellites hallii sp. nov. 


Hexacarpellites hallii sp. nov. 
Plate 51, figures 1—4 


Etymology.— This species is named after William 
and Gertrude Hall of Vancouver, Washington recog- 
nizing their encouragement and support of the Clarno 
paleobotany project. 

Description.— Fruit subglobose, prolate, with six ra- 
dially connate carpels, base and apex rounded, surface 
texture unknown; length 5.1 mm, equatorial diameter 
4.3 mm; with six vertically elongate locules up to 4.5 
mm high, 0.8 mm wide and 1.1 mm dorsiventral thick- 
ness; septa thick, with obvious suture lines; one seed 
per locule, placentation axile; seeds shrunken within 
locules, seed coat composed ofa thin layer of polygonal 
to spherical, tan cells, 20-35 um in diameter; apparent 
embryo within seed 0.41 mm thick dorsiventrally, O.1 
mm wide, preserved in dark brown silica; central axis 
of fruit thick and prominent from base to apex, filled 
with dark fibers and vascular tissue. 

Specimen.— Holotype: USNM 424653. 


102 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Discussion. — Hexacarpellites hallii is represented by 
a single specimen. Although it appears to have a rough 
outer surface (Pl. 51, fig. 1), the exterior of the fruit is 
fragmented away, possibly having adhered to the sed- 
imentary matrix from which it was collected, leaving 
no evidence of original surface detail. Serial sectioning, 
however, revealed very good internal preservation, in- 
cluding a seeds and embryo within each locule (Pl. 51, 
figs. 2-4). Successive transverse sections showed that 
the locules are axially elongate and single-seeded. The 
distal margin of each locule cast is exposed in a median 
longitudinal groove of the fruit. This may be an artifact 
of the way that the fossil was removed from the matrix, 
but if it is an original feature of the fruit it suggests 
that dehiscence was loculicidal. 


Genus JOEJONESIA gen. nov. 


Etymology.—Named after Joseph Jones III, director 
of Hancock Field Station, Fossil, Oregon, recognizing 
his support and encouragement of paleobotanical field 
studies at the Nut Beds. 

Type species.—Joejonesia globosa sp. nov. 


Joejonesia globosa sp. nov. 
Plate 51, figures 5-8 


Etymology.— Globus (L = ball), referring to the fruit 
shape. 

Description.—Fruit globose, unilocular, bivalved, 
height 8 mm (est.), equatorial diameter 8.0 mm, with 
a reticulum of thin veins over the surface; endocarp 
wall 1.3 mm thick, composed of isodiametric suban- 
gular sclereids typically 90-150 um in diameter, small- 
er toward the locule (40-60 um), locule lining unise- 
riate, composed of poorly preserved rectangular cells, 
40 um thick; locule smooth, with a keel in the plane 
of dehiscence. 

Specimen.— Holotype: UF 9260. 

Discussion.— This species is represented by a single 
fragmentary specimen. It is sufficiently complete, how- 
ever, to discern both internal and external morphology 
of the endocarp, and the anatomical preservation is 
excellent. The salient features of Joejonesia are globose 
shape, reticulate surface, bivalved endocarp (Pl. 51, 
fig. 5), the smooth, keeled locule (Pl. 51, figs. 6, 7) and 
the large sclereids composing the endocarp (PI. 51, fig. 
8). Although the locule in this specimen is empty, its 
form suggests that there was only a single seed. In size 
and bivalvate construction, it brings to mind Prunus, 
but Prunus has smaller sclereids composing the wall 
and the dehiscence plane is developed on only one side 
of the endocarp (cf. Pl. 35, figs. 11-15). 


Genus LIGNICARPUS gen. nov. 


Etymology.— Lignum (Gr = wood) + karpos (Gr = 
fruit). 


Type species.— Lignicarpus crassimuri sp. nov. 


Lignicarpus crassimuri sp. nov. 
Plate 51, figures 9-16 


Etymology.— Crassi (L = thick) + murus (L = wall). 

Description.—Endocarp ovoid, elliptical to circular 
in cross section, unilocular, single-seeded, base and 
apex obtuse-rounded, length 9.7-11.4 mm, maximum 
width 6.0-6.1 mm, minimum width 3.7 mm; endocarp 
surface rough-textured, with longitudinally striate fi- 
brous ribs, endocarp wall 1.4-2.0 mm thick, composed 
of longitudinally oriented fibers 50-88 um diameter, 
with evenly distributed clusters (as seen in cross 5ес- 
tion) of smaller vertically oriented vascular strands; 
locule narrow-pyriform, rounded basally, tapering to 
a point apically. Seed pendulous?, conforming in shape 
to the locule, with a thin seed coat. 


Specimens.—Holotype: UF 6111. Paratypes: UF 


6578, 9765. 

Discussion.—These thick-walled unilocular endo- 
carps bear a superficial resemblance to those of Nyssa. 
However the fibers of Lignicarpus crassimuri run vet- 
tically, unlike those of Nyssa which form swirling tracts 
of varying orientation (cf. Pl. 11, fig. 19), and Nyssa 
does not have vascular bundles scattered through the 
endocarp. 


Genus LIGNIGLOBUS gen. nov. 
Etymology.—Lignum (L = wood) + globus (L = 


ball), referring to the woodiness and globose shape of 


the endocarp. 
Type species.— Ligniglobus sinuosifibrae sp. nov. 


Ligniglobus sinuosifibrae sp. nov. 
Plate 53, figures 1-8 


Etymology.— Sinuosus (L = full of bendings) + fibra 
(L = thread, fiber), referring to the sinuous arrangement 
of fibers over the surface of the endocarp. 

Description.— Fruit subglobose, unilocular with а 
protrusion at one end, length 20.0-30.5 mm, equato- 
rial diameter 20.0-25.1 mm; surface verrucate with 
fine, wavy, longitudinal striations; pericarp 4.2-6.3 mm 
thick composed successively of: 1) locule lining 120 
um thick, cells not well preserved, 2) main inner layer 
3.7 mm thick of anticlinally oriented septate fibers 17- 
25 um thick radiating outward from locule; 3) a 1922 
mm-thick zone of isodiametric parenchyma cells 50- 
80 um in diameter; 4) an outer layer approximately 1 
mm thick with periclinally and horizontally oriented 
fibers; 5) epicarp composed of periclinal files of rect- 
angular cells 35-60 um wide, 12-30 um thick, appat* 
ently with ducts or vasculature running through this 
layer; locule globose to elongate. 

Specimens.—Holotype: UF 8483. Paratypes: UF 
8480, 9894, USNM 312752 (Bones, 1979, pl. 1, fig- 
7), 354386. 


| 
| 
| 
| 
| 
| 
| 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 103 


Discussion.— Ligniglobus is represented by permin- 
eralized fruits that reveal woody internal structure in 
sections and fractured surfaces. Because ofits relatively 
large size, thick wall, and single locule (P1. 53, figs. 1- 
4), this fruit type is superficially similar to Scabrae- 
carpium clarnense (cf. Pl. 59, figs. 1-7) but is distin- 
guished by its more globose shape and finer, narrower 
endocarp fibers and by the lack of fibrovascular bun- 
dles in the endocarp. 


Genus LUNATICARPA gen. nov. 


Etymology.— Lunat (L = crescent moon) + karpos 
(Gr = fruit), referring to the crescent shaped outline of 
the fruit. 

Type species.— Lunaticarpa curvistriata sp. nov. 


Lunaticarpa curvistriata sp. nov. 
Plate 52, figures 1—12 


Etymology.— Curvus (L = bent) + striata (L = line), 
referring to the endocarp surface pattern. 

Description.—Endocarp reniform, bilaterally sym- 
metrical, laterally compressed in the plane of sym- 
metry, unilocular, single-seeded; dorsal surface con- 
vex-rounded, with a rounded keel in the plane of 
Symmetry, ventral side convex-rounded to nearly 
straight with a circular, centrally positioned funicular 
Opening; dorsiventral length 2.6-4.9, avg. 3.7 mm 
(SD=0.51, n=32), width 3.2-5.0, avg. 4.1 mm, 
(SD=0.53, n=32), thickness 1.0-2.2, avg. 1.9 mm 
(SD=0.35, n=32); surface finely striate with the stri- 
ations on the lateral faces arched parallel to the cur- 
Vature of the dorsal surface; fruit surface following con- 
tour of the curved seed, and the bulging median condyle; 
Pericarp 200-250 um thick around the dorsal side, 
about five times thicker adjacent to the funicular open- 
Ing, composed of: an inner layer 110 um thick of iso- 
diametric cells 12.5-30.0 um in diameter and an outer 
layer 100 um thick composed of fibers oriented parallel 
to the surface. Locule cast crescent-shaped to nearly 
Straight, with a sharp median keel on the dorsal side 
and a ventral funicular plug, smooth to very faintly 
transversely ruminate, one of the limbs rounded, the 
Other with a pointed tip. Seed curved, conforming to 
the shape of the locule. 

Specimens.— Holotype: USNM 435020. Paratypes: 
UCMP 10717-10719, UF 6336-6341, 8582, 8767, 
8773, USNM 355653, 422527, 422528, 435018, 
435019, 435021, 435022, 435024-435037, 435045, 
435046. 

Discussion. — This species is represented by permi- 
Deralized endocarps and numerous isolated locule casts. 
Distinctive endocarp features of Lunaticarpa are the 
teniform shape, dorsal keel, curved striations on the 
lateral faces (Pl. 52, figs. 1-3), and the large condylar 
Cavity about which the locule is curved (Pl. 52, figs. 


10, 11). The curvature and faint transverse rumina- 
tions of the locule cast bring to mind Menispermaceae. 
Similar, unornamented endocarps occur in extant Pa- 
chygone. However, this similarity may be the result of 
convergence; I have not been able to confirm affinities 
with Menispermaceae. 


Genus MICROPHALLUS gen. nov. 


Etymology.— Micro (Gr = small) + phallus (Gr), 
referring to the penial appearance of these specimens. 
Type species. — Microphallus perplexus sp. nov. 


Microphallus perplexus sp. nov. 
Plate 53, figures 9-17 


Etymology. — The epithet perplexus refers to the baf- 
fling affinities of this taxon. 

Description. —Seed pyriform or tear-drop shaped in 
outline, bilaterally symmetrical, dorsiventrally com- 
pressed with a central depression (chalaza?) on the ven- 
tral surface, and a raised circular area (hilum?) on the 
dorsal surface; length of complete seed 4.8—6.7, avg. 
5.5 mm (SD=0.45, n=7), width 3.2-4.7, avg. 3.8 mm, 
(SD=0.50, п=7), thickness 2.3-3.2, avg. 2.7 mm 
(SD=0.34, n=6); seed coat thick (0.6 mm) comprised 
ofa uniseriate layer of elongate thick walled cells which 
form coarse surface striations that radiate from the 
periphery of the dorsal raised area and the from the 
center of the ventral face. Embryo forming a circular 
loop in the main seed body, and a pronounced apical 
extension (radicle) about 0.7 mm in diameter and up 
to 2 mm long that protrudes through the apex of the 
seed; length of embryo cast 4.4-6.3, avg. 5.5 mm 
(SD=0.72, n=5), width 2.5-3.2, avg. 2.8 mm (SD=0.27, 
n=5), thickness 1.2-1.6, avg. 1.4 mm (SD=0.15, n=5). 

Specimens.—Holotype: UF 5243, complete seed. 
Paratypes: UF 6346, 9280, USNM 355414, 355391, 
355685 (2 specimens), complete seeds; OMSI Pb1839, 
UCMP 10739, 10740, UF 5242, 9281, 9282, USNM 
355685 (3 specimens), 434965, embryo casts. 

Discussion.—Seeds of Microphallus present a dis- 
tinctive morphology and are easily recognized because 
the embryo, usually preserved in white chalcedony, 
resembles a screw eye (Pl. 53, fig. 13). The loop in the 
embryo cast may represent curved cotyledons, but in 
most specimens the loop is closed with no indication 
of where the cotyledons might terminate. I interpret 
the raised dorsal area on the seed (Pl. 53, figs. 10-12, 
15, 16) to be the hilum and the ventral cavity (Pl. 53, 
figs. 9, 14) to be the chalaza, but as yet, I have no clue 
as to the systematic affinities of this fossil. 


Genus NEPHROSEMEN gen. nov. 


Etymology.— Nephro (Gr = kidney) + semen (L = 
seed) referring to the roughly kidney-shaped outline of 
the seed. 


104 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Type species.— Nephrosemen reticulatus sp. nov. 


Nephrosemen reticulatus sp. nov. 
Plate 54, figures 6-13 


Etymology.—reticulatus (L = netlike), referring to 
the seed surface pattern. 

Description.—Seed reniform, bilaterally symmetri- 
cal, laterally compressed in the plane of symmetry, 
more or less rounded in lateral profile, with a concave 
notch at the hilar end, micropyle pointed, situated ad- 
jacent to hilum; seed height (hilum to dorsal edge) 2.2- 
3.1, avg. 2.7 mm (SD=0.28, n=19), width 2.2-3.0, avg. 
2.5 mm (SD=0.23, n=19), thickness 1.1-1.9, avg. 1.3 
mm (SD=0.20, n=19); smoothly contoured, surface 
covered by a fine reticulum of pentagonal and hexag- 
onal cells 49-62 um in diameter with thick anticlinal 
walls, these cells aligned in rows radiating from the 
micropyle area. 

Specimens.—Holotype: UF 6500. Paratypes: UF 
6389, 6573, 9167, 9376-9382, USNM 355047, 355052, 
355403, 424683424686, 424746-424749. 

Discussion.— Nephrosemen is readily recognized in 
the Nut Beds collections by the kidney-shaped lateral 
profile, lateral compression and reticulate surface. 
Specimens vary from longer than wide (Pl. 54, figs. 6- 
9, to wider than long (Pl. 54, fig. 11). The closely ad- 
jacent hilum and micropyle (PI. 54, fig. 7) suggest that 
the seed was anatropous. The seed shape and reticulate 
surface bring to mind seeds of extant Theaceae. They 
are much larger than the seeds of Cleyera (cf. Pl. 38, 
figs. 1-5), yet the surface reticulum (Pl. 51, fig. 10) is 
much finer, and a curved embryo has not been detected 
in Nephrosemen. There is an even closer resemblance 
to seeds of genera in the Solanaceae, for example, Man- 
dragora, Solanum (part), Physocarpus, in the shape 
and reticulate surface of the seed. Whether the embryo 
is curved or coiled in the fossil, as in Solanaceae, re- 
mains undetermined. 


Genus OMSICARPIUM gen. nov. 


Etymology.—This fruit genus is named for OMSI, 
the Oregon Museum of Science and Industry, recog- 
nizing the role of this museum in supporting this pro- 
ject: 

Type species.— Omsicarpium striatum sp. nov. 


` Omsicarpium striatum sp. nov. 
Plate 52, figures 13-16 


Etymology.— Stria (L — furrow, line) referring to the 
striated endocarp surface. 

Description.— Fruit pyriform, more or less circular 
in cross section, base and apex rounded, unilocular; 
height 33.5 mm, width 20.0 mm; ornamented with 
prominent, sinuous, closely spaced longitudinal stria- 


tions, pericarp 2.4 mm thick, with an apparent lon- 
gitudinal suture on one side, anatomy and seed struc- 
ture not preserved. 

Specimen.— Holotype: OMSI Pb265. 

Discussion.— This specimen is preserved in calcite, 
without cellular details. The surface detail is well pre- 
served, and the specimen is clearly unilocular with a 
relatively thick wall. In general form it is similar to 
Pruniticarpa. 


Genus PASTERNACKIA gen. nov. 


Etymology.— Named for Ellen Pasternack recogniz- 
ing her assistance in collecting at the Nut Beds. 
Type species. — Pasternackia pusilla sp. nov. 


Pasternackia pusilla sp. nov. 
Plate 55, figures 1—4 


Etymology.— The epithet pusilla (L — very little) re- 
fers to the small size of these seeds. 

Description. —Seed anatropous, ovoid, bilaterally 
symmetrical, nearly circular in transverse section, ta- 
pering gradually to the acute hilar/micropylar end, 
rounded-obtuse at the opposite end; length 1.1-1.2, 
avg. 1.1 mm (SD=0.05, n=5), width 0.7-0.8, avg. 0.74 
mm (SD=0.05, n=5); surface reticulate, formed by 16 
to 22 meridionally aligned rows of cells with thickened 
anticlinal walls, these cells four- to five-sided in surface 
view; the rows ofcells along the raphal side (?) narrower 
than those over the rest of the seed surface. 

Specimens.— Holotype: UF 9604. Paratypes: UF 
9605-9608. 

Discussion.— Pasternackia seeds are preserved as 
small ovoid chalcedony casts with a distinctive retic- 
ulate surface formed by large, meridionally aligned seed 
coat cells (Pl. 55, figs. 1-4). There is a resemblance to 
seeds of Saurauria in the Actinidiaceae, but the me- 
ridional alignment of cells is more strongly asymmet- 
rical. The hilar end is damaged in each of the fossils 
so that the precise morphology of the hilum and mi- 
cropyle could not be determined. 


Genus PILEOSPERMA gen. nov. 


Etymology.— Pileos (Gr = cap) + spermum (Gr = 
seed) 
Type species. — Pileosperma minutum sp. nov. 


Pileosperma minutum sp. nov. 
Plate 55, figures 5-8 


Description.—Seed pyriform, bilaterally symmetri- 
cal, rounded basally and laterally, seed broadest near 
the base with the wall tapering gradually toward the 
pointed micropylar end, abruptly more tapered near 
the apex to form a broad-conical tip, seed length 2.5- 


3.2, avg. 2.7 mm (SD=0.25, n=6), width in plane of 


| 
| 
| 
| 
| 
| 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 105 


Symmetry 1.2-1.8, avg. 1.5 mm (SD=0.20, n=6), width 
at right angles to plane of symmetry 1.0-1.4, avg. 1.3 
mm (SD=0.15, n=6); with a raised circular cap (cha- 
laza?) at the base facing ventrally, surface microreti- 
Culate, made up of isodiametric polygonal cells 15-20 
km in diameter, sometimes with faintly superimposed 
Impression of large meridionally arranged rectangular 
Cells. 

Specimens.—Holotype: UF 6606. Paratypes: UF 
6425, 9588-9591; USNM 424784424787. 

Discussion.—These small seeds are preserved as 
Chalcedony casts with well preserved cell outlines on 
the surface (Pl. 55, fig. 7). The raised circular basal cap 
55, figs. 5, 8) that may represent the chalaza is a 
Conspicuous feature of this fossil genus. This cap is 
Sometimes detached, but is still evident because of the 
Scar left behind (Pl. 55, fig. 6). There is no obvious 
Taphe, but there commonly is a broad, shallow ventral 
groove leading between the base and apex. 


Pileosperma ovatum sp. nov. 
Plate 55, figures 9-11 


Description. —Seed pyriform, bilaterally symmetri- 
Cal, pointed at the apical micropyle, rounded basally 
апа laterally, broadest near the base, gradually tapered 
all the way to the apex, length 2.0-3.0, avg. 2.5 mm 
(SD—0.32, n=32), broadest width 1.5-2.2, avg. 1.9 mm 
(SD—0.19, n=33), narrowest width 1.3-2.0, avg. 1.6 
mm (SD=0.16, n=33), with a raised circular cap (cha- 
laza?) at the base. Surface microreticulate, made up of 
ISodiametric polygonal cells 23-35 um in diameter, 
Sometimes with faintly superimposed impression of 
arge meridionally arranged rectangular cells. 

Specimens.— Holotype: USNM 424751. Paratypes: 
UF 6420, 6422-6423, 6499, 9722, 9723. USNM 
424726, 424755, 424758, 424760, 424762, 424764, 
424766, 424767, 424769-424783, 424787. 

Discussion.— Although clearly congeneric with Pi- 
€osperma minutum, P. ovatum is distinguished by its 

Toader shape, and by the uniform curvature of the 
lateral walls toward the apical tip. 


Genus PISTACHIOIDES gen. nov. 


Etymology.— Pistachia + oides (Gr = like, resem- 
ling), indicating the similarity to the common pista- 
Chio nut, 
Type species.— Pistachioides striata sp. nov. 


Pistachioides striata sp. nov. 
Plate 54, figures 1—5 


Etymology.— Stria (L = furrow, line), referring to 
Ше striated surface of the endocarp. 
{ Description. —Fruit ovate in face view, nearly ellip- 
tical in lateral view, rounded-quadrangular in cross 


section, base rounded, apex pointed, bilaterally sym- 
metrical, unilocular, length 21-24 mm, width across 
plane of symmetry 10.7-13.6 mm, dorsiventral thick- 
ness 8.7-14.2 mm, with a keel in the plane ofsymmetry 
showing a suture line along which two valves were 
probably joined; endocarp wall thin (0.4 mm), com- 
posed of diagonally oriented fibers, attachment scar 
elliptical, elongate in the plane of symmetry. 

Specimen.— Holotype: USNM 422387. Paratype: UF 
6440. 

Discussion.— Pistachioides is represented by two en- 
docarp casts from the Nut Beds. The holotype (Pl. 54, 
figs. 1-5) is replaced by chalcedony; the paratype is 
replaced by calcite. Characters of shape, size, keel and 
dehiscence plane coincident with the plane of sym- 
metry, diagonally oriented surface striations, and the 
attachment scar which is elongate in the plane of sym- 
metry, correspond closely to those of Pistacia vera, the 
commercial pistachio nut. The basal margin, as viewed 
laterally in the plane of symmetry, is oblique, as it is 
in Pistachia. I have hesitated to place it within this 
extant anacardiaceous genus because of the lack of cor- 
roborating anatomical details in the fossils. Internal 
anatomy is not preserved in the fossil specimens; the 
fossils, however, appear to have a fibrous outer layer, 
whereas the outer layer in Pistacia is formed by more 
or less isodiametric sclereids. 


Genus POLLOSTOSPERMA gen. nov. 


Etymology.— РоПочох (Gr = smallest, least) + sper- 
ma (Gr = seed), because these seeds are among the 
smallest from the Nut Beds. 

Туре species.— Pollostosperma dictyum sp. nov. 


Pollostosperma dictyum sp. nov. 
Plate 56, figures 9-12 


Etymology. — Diktyon (Gr = net) referring to the re- 
ticulate surface of the seed. 

Description.—Seed laterally compressed, wide-ovate 
in lateral view, lensoidal in transverse section, bilat- 
erally symmetrical, length 1.1-1.3 mm, width 0.9-1.2 
mm, thickness across plane of symmetry 0.6-0.9 mm, 
with a keel in the plane of symmetry, rounded basally 
and pointed apically as viewed laterally; surface retic- 
ulate, with cells ca. 50 um in diameter.” 

Specimens.—Holotype: UF 9830. Paratypes: UF 
9831-9840, USNM 355448. 

Discussion. — This species is one of the more abun- 
dant seed types at the Nut Beds although it is easily 
overlooked in the field because of its small size. In 
many specimens the reticulate surface is damaged in 
the process of removal from the matrix. In sorting 
through numerous seeds of this type collected by Tho- 
mas Bones, however, a few specimens with well pre- 


106 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


served external surface were found (e.g., Pl. 56, figs. 
9-12). The affinities are still unknown. 


Genus POLYGRANA gen. nov. 


Etymology.— Poly (Gr = Many) + granum (Gr = 
seed), referring to the large number of seeds within this 
fruit type. 

Type species. — Polygrana nutbedense sp. nov. 


Polygrana nutbedense sp. nov. 
Plate 56, figures 1-8 


Etymology.— The epithet refers to the Nut Beds lo- 
cality. 

Description.— Fruit a subovoid, prolate, multiseed- 
ed berry with slight irregular depressions and bulges, 
roughly circular in cross section, length 9.1-10.1 mm, 
equatorial diameter 6.1-8.4 mm, perianth hypogy- 
nous; scar forming a persistent circular flange at the 
fruit base; fruit surface smooth, with occasional fine, 
unbranched longitudinal fibrovascular bundles; uni- 
locular with many seeds; fruit wall ca. 0.4 mm thick, 
placentation apparently parietal, with seeds embedded 
in a tissue of parenchyma cells 50-240 um in diameter, 
and/or with groups of narrow parallel fibers. Seeds 
ovoid, 1.0 mm long by ca. 0.5 mm їп equatorial di- 
ameter, orthotropous (?), seed coat 40-60 um thick 
with a conspicuous layer of about eight rows of anti- 
clinally flattened polygonal cells, embryo straight. 

Specimens.— Holotype: USNM 355493. Paratypes: 
UF 5665, 5673. 

Discussion.— Polygrana is known only from a few 
permineralized multiseeded berries. The thin exocarp 
appears to have been leathery, and the mesocarp pulpy, 
resulting in distorted, somewhat squashed specimens 
(e.g., Pl. 56, fig. 6). The placentation apparently is pa- 
rietal, although many of the seeds appear to be floating 
within parenchymatous tissue some distance from the 
margin (Pl. 56, figs. 5, 6). The number of carpels is 
uncertain. The fruits bear a resemblance to certain Fla- 
courtiaceae, including Banara. I have not found a 
modern genus with a seed coat of conspicuously flat- 
tened polygonal cells like that of this fossil (Pl. 56, fig. 
8). 


Genus PRUNITICARPA gen. nov. 


Etymology.—Prunus + пе (L, Gr = having Ше na- 
ture of) + karpos (Gr = fruit), referring to the general 
resemblance to extant Prunus. 

Type species.— Pruniticarpa cevallosii sp. nov. 


Pruniticarpa cevallosii sp. nov. 
Plate 56, figures 13-16; Plate 57, figure 20 


Etymology.— The epithet cevallosii is named in гес- 
ognition of the contributions of Sergio Cevallos-Ferriz 
to fossil wood and fruit research. 


Description.—Endocarp subpyriform, nearly circu- 
lar in cross section, pointed apically and basally, bi- 
laterally symmetrical, unilocular, single-seeded, length 
39 mm, width 24 x 23 mm, surface rough, with a 
single low, rounded longitudinal ridge running from 
base to apex at the position of carpel closure in the 
plane of symmetry, endocarp wall 3.8-4.3 mm thick, 
composed of compact sclereids, grading from an inner 
layer of small isodiametric sclereids 20-38 um in di- 
ameter through a middle layer with anticlinally elon- 
gate cells 100-150 um long and 25-65 um wide, to the 
outer part with large rounded isodiametric cells 37-75 
um in diameter; locule lining loosely adherent to the 
endocarp, composed of a three- to five-seriate layer of 
small globose cells, 20-30 um in diameter, with a uni- 
seriate layer of scattered rectangular peg-like cells 40- 
50 um in diameter. Locule cast rounded basally, point- 


ed apically with a longitudinal keel corresponding 10. 


the dehiscence line, 25.7 mm long, 7.7 mm wide. 

Specimens.— Holotype: UF 9209. Paratypes: USNM 
355194, 446087. 

Discussion.— This large unilocular endocarp is known 
from only a few well preserved specimens. The holo- 
type was sectioned transversely, showing the thick en- 
docarp wall, nearly circular outline of the endocarp 
and locule and plane of dehiscence (Pl. 56, fig. 15) 
Although it resembles Prunus in having a thick-walled, 
woody, unilocular endocarp and plane of dehiscence 
running longitudinally on only one side of the fruit, I 
have not observed any extant species of Prunus with 
endocarps that are circular in cross section. 


Genus PTERONEPELYS gen. nov. 


Etymology.— Pteron (Gr = wing) + epelys (Gr = 
stranger). 
Type species.— Pteronepelys wehrii sp. nov. 


Pteronepelys wehrii sp. nov. 
Plate 57, figures 1-3; Text-figure 20 


Etymology.— The species is named in honor of Wes- 
ley Wehr, who has provided excellent comparative ma- 
terial of this taxon from the Republic flora, Washing- 
ton. 

Description. —Samara consisting of a laterally flat- 
tened, ovoid endocarp with an elongate, more or less 
elliptical wing; samara length 10.0-13.9 mm, width 
3.7-4.3 mm; endocarp length 3.5-4.5 mm, width 2.6- 
3.0 mm, thickness ca. 2-2.5 mm; wing with a straight 
to slightly convex upper margin and a convex lower 
margin joined distally in an acute angle, three long!” 
tudinal veins arising from the nutlet, joining at the 
distal tip of the wing: a thick vein closely following the 
upper margin of the wing, another thick vein situate 
between the midline of the wing and the upper wing 
margin, and a thin vein following the lower marg” 


| 
| 
| 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 107 


Situated between the lower margin and midline of the 
Wing; apex of nutlet, opposite Ше wing attachment, 
giving rise to four (or five?) stout sepals 1.5 mm high 
and 1.5 mm wide with rounded apices, and two central 
Style arms. 

Specimens.— Holotype: OMSI Pb1018. Paratypes: 
OMSI Pb1381, Nut Beds; UF 11696, Clarno White 
Cliffs locality; UWBM 36963, 36966, 36976, 36977, 
54650, Republic flora. 

Discussion.— This species is represented in the Nut 
Beds by a few specimens preserved as impressions in 
Siltstone from the leaf layer of Face 3 (Pl. 57, figs. 1, 
2). Although it has not been described previously, the 
Same species is also known from the Clarno White 
Cliffs locality (Pl. 57, fig. 3) and from the Middle Eo- 
Сепе Republic flora of northeastern Washington (Wolfe 
and Wehr, 1987). Details of the sepals and styles are 
Em clearly preserved in the specimens from Repub- 
ic, 

Superficially, Pteronepelys fruits resemble those of 
Cedrelospermum (cf. Pl. 43, figs. 16-18); however, 
Pteronepelys is distinguished by fewer veins in the wing, 
by the lack of a secondary wing, and the position of 
the perianth relative to the wing. The sepals protrude 
from the opposite side of the nutlet from the wing. 
From the orientation of the styles and perianth, it ap- 
Pears that the pedicel attachment was at the other end 
Of the fruit, at the tip of the wing (Text-fig. 20). 


Genus PULVINISPERMA gen. nov. 


Etymology.—Pulvinis (L = cushion, pad, pillow) + 
Spermum (Gr = seed). 
Type species.— Pulvinisperma minutum sp. nov. 


Pulvinisperma minutum sp. nov. 
Plate 57, figures 4-8 


Etymology. — Minutus (L = small, little), referring to 
the seed size. 

Description.—Seed more or less pillow-shaped, bi- 
laterally symmetrical, laterally compressed with a keel 
Ш the plane of symmetry, rounded-rhomboidal in lat- 
ега] view, more or less lensoidal in face view and in 
Cross section, length 0.9-1.3, avg. 1.1 mm (SD=0.13, 
1—10), width in plane of symmetry 0.7-0.8, avg. 0.76 
mm (SD=0.05, n=10), thickness across the plane of 
Symmetry 0.5-0.7, avg. 0.56 mm (SD=0.07, n=10), 
Tounded basally and apically, hilum and elliptical scar 
Ш the plane of symmetry at the junction of the apical 
4nd ventral margins; pointed, micropylar projection 
Оп the opposite side from the hilum at the junction of 
the apical and dorsal margins. 

Specimens.—Holotype: UF 9740. Paratype: UF 
9741-9743, 9841-9850. 

Discussion.— This species is represented by numer- 
Ous small chalcedony seed casts. Salient features in- 


lem 


Text-figure 20.— Pteronepyles wehrii gen. et sp. nov. 


clude the rhomboidal outline in lateral view and keel 
in the plane of symmetry. The affinities remain un- 
known. 


Genus PYRISEMEN gen. nov. 


Etymology.— Pyrum (L = pear) + semen (Г. = seed), 
referring to the pear-like shape of the seed. 
Type species.— Pyrisemen attenuatum sp. nov. 


Pyrisemen attenuatum sp. nov. 
Plate 57, figures 9-12 


Etymology.— Attenuatus (L = drawn out, tapered), 
referring to the narrowed seed apex. 

Description.—Seed/achene pyriform, asymmetrical, 
attenuate-pointed apically, rounded basally, laterally 
flattened, elliptical in cross section, length 2.4—4.3, avg. 
3.1 mm (SD=0.58, n=9), width 1.2-3.5, avg. 1.3 mm 
(SD=0.91, n=9), thickness 1.8-2.3, avg. 2.1 mm 
(SD=0.19, n=9); base with a recessed circular scar, 0.8— 
1.0 mm in diameter, surface smooth. Inner body sim- 
ilar in shape to outer surface, forming a silhouette in 
translucent specimens. 

Specimens.— Holotype: UF 6463. Paratypes: UCMP 
10626-10629, UF 6400-6404, 6464-6466, USNM 
355373, 355419 (6 specimens), 435049, 435059. 

Discussion.—This species is usually preserved in 
white or translucent chalcedony. Although internal cell 
structure has not been preserved, some specimens show 
the silhouette of an internal seed coat in transmitted 
light (Pl. 57, figs. 11, 12). Possibly they are achenes 
tapering apically toward the style with the circular scar 
representing the pedicel attachment. Alternatively, they 
may represent bitegmic seeds with a circular basal hi- 
lum tapering toward the micropyle. Seeds that are at 
least superficially similar occur in the Flacourtiaceae, 
e.g., Oncoba. 


Genus QUINTACAVA gen. nov. 
Etymology.— Quinta (L = five) + cavus (L = hollow, 
hole), referring to the five prominent locules of the 
fruit. 
Type species.— Quintacava velosida sp. nov. 


108 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Quintacava velosida sp. nov. 
Plate 57, figures 13-19; Text-figure 21 


Etymology.— Velo (L = cover, conceal) + sidus (L 
= star) referring to the star-shaped locule arrangement 
as seen in transverse section. 

Description.— Fruit pentacarpellate, rounded-ob- 
conical, broadest basally, base more or less flat, with 
a circular pedicel scar in the center, apex rounded; 
length 6.9 mm, width 7.0-7.8 mm; central axis vas- 
cularized by a thin strand; five radially arranged, slen- 
der, elongate locules that are lanceolate in cross section 
alternating with large lacunae that are more or less 
triangular in cross section; lacunae occupying up to 
five times as much area as the locules, each locule 
separated from the adjacent lacuna by a thin wall, 0.15 
mm thick, composed of five to twelve rows of axially 
oriented fibers 6-10 um in diameter with scattered 
vertical resin or gum ducts, these ducts circular in cross 
section, 12-25 um in diameter; locule lining composed 
of a uniseriate layer of fibers varying in orientation 
from nearly horizontal near the fruit axis to nearly 
vertical toward the periphery of the fruit; outer wall of 
fruit 0.56 mm thick composed of isodiametric paren- 
chyma cells, 40-60 um in diameter; seeds not ob- 
served. 

Specimen.— Holotype: USNM 424805. 

Discussion.— This species is represented by a single 
well-preserved specimen. In cross section the arrange- 
ment of locules resembles a five-rayed star (Pl. 57, fig. 
16). The anatomy of the septa, with vertically oriented 
fibers and conspicuous longitudinal ducts (Pl. 57, figs. 
17, 18) are distinctive for this species. The outer wall 
of the fruit was soft or spongy, made up of parenchyma 
(Pl. 57, fig. 19) and the hard, fibrous wall around each 
of the locules appears to have functioned as a pyrene 
with dorsal dehiscence; indeed, two of the locules have 
split along their midline on the distal side (Pl. 57, fig. 
16). The number and arrangement of locules is con- 
sistent with fruits of the Maloideae subfamily of the 
Rosaceae (Rohrer et al., 1991); however, the affinities, 
whether in the Rosaceae or elsewhere, remain to be 
determined. 


Genus SAMBUCUSPERMITES gen. nov. 


Etymology.—Sambucus + spermum (Gr = seed) + 
ites (Gr = like, having the nature of). 
Type species. — Sambucuspermites rugulosus sp. nov. 


Sambucuspermites rugulosus sp. nov. 
Plate 58, figures 10-14 


Description. —Seed small, elliptical to obovate in face 
view, slightly tapered toward the hilum, rounded at 
the opposite end, dorsiventrally flattened, sometimes 
concave ventrally and convex dorsally, length 1.2-1.5, 


avg. 1.4 mm (SD=0.12, n=5), width 0.9-1.2, avg. 1.0 
mm (SD=0.11, n=5), thickness 0.4-0.5, avg. 0.46 mm 
(SD=0.05, n=5); surface with irregular, wavy horizon- 
tal rugulae, six to seven rugulae from base to apex. 

Specimens.— Holotype: UF 6424. Paratypes: OMSI 
PB1716 (3 specimens), UF 6396-6399, 6426-6428, 
USNM 353980 (4 specimens). 

Discussion.—This species is preserved as small si- 
licified seeds, readily recognized by the ovate to ellip- 
tical shape, prominent horizontal rugulae (Pl. 58, figs. 
10-14) and dorsiventral compression (Pl. 58, fig. 14). 
The seeds resemble those of modern Sambucus in gen- 
eral morphology and outline of surface, including the 
rugulate sculpture and the small circular aperture at 
the hilum. Sambucuspermites differs from extant spe- 
cies of Sambucus, however, in its small size (modern 
species are about 3 mm long), and fewer rugulae. Seeds 
of the Recent species S. nigra L., S. ebulus L., S. puch- | 
ella, with which the fossil was compared, are more 
elongate in outline and have more rugulations than the 
fossil. Internal anatomy ofthe fossils was not observed. 
The name Sambucuspermites indicates the morpho- 
logical similarity to Sambucus seeds, but does not nec- 
essarily imply a close relationship. 


Genus SCABRAECARPIUM gen. nov. 


Etymology.— Scabra (L = rough, scabby) + karpos 
(Gr — fruit). 
Type species.— Scabraecarpium clarnense sp. nov. 


Scabraecarpium clarnense sp. nov. 
Plate 59, figures 1-7 


Etymology.— The epithet refers to the Clarno For- 
mation. 

Description.— Fruit ovoid to obovoid, circular to el- 
liptical in cross section, unilocular, base and арех 
rounded to obtuse-pointed, sometimes with a large 
circular recessed scar at base, fruit length 32.0-43.5, 
avg. 37.3 mm (SD=3.92, n=7), minimum width 23.0- 
28.5, avg. 26.3 mm (SD=2.19, n=8), maximum width 
26.4—34.5 avg. 31.4 mm (SD=4.36, n=8), surface warty» 
woody to abraded; wall thick (4.5-10.5 mm) consisting 
of three layers: 1) an inner layer of radially arranged 
files of anticlinally elongate rectangular thin-walled cells 
15-50 um wide, 50—250 um high, successively wider 
cells occur distally, 2) middle layer of thicker-walled 
cells, also anticlinally elongate and arranged in radial 
files, cells rectangular, 30-75 um wide, 50-150 um 
high, 3) outer layer of isodiametric parenchyma cells 
50—70 um in diameter; fibrovascular bundles 300-400 
um in diameter common, running obliquely with sin- 
uous course through the middle layer and vertically 
through the outer parenchyma layer; locule circular 10 
elliptical in cross section, 8—11 mm in diameter. 


| 
| 
| 
| 
| 
| 
| 
| 
| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 109 


5mm 


Text-figure 21.— Quintacava velosida gen. et sp. nov., fruit in trans- 
Verse section showing five locules, large lacunae, and relatively thick 
Outer wall. 


Specimens.— Holotype: USNM 355066. Paratypes: 
UF 8481, 8482, USNM 354200, 354201, 354385, 
354421, 354667, 422397, 424811. 

Discussion.— Scabraecarpium is a thick-walled uni- 
locular fruit type that is among the larger fruit types 
from the Nut Beds. The fruit wall is usually permi- 
Neralized, and appears to have had a woody texture. 
Specimens vary from circular (Pl. 59, figs. 2, 5) to 
elliptical (Pl. 59, figs. 4, 6) in cross section. The fruit 
Wall is similar in the anticlinal arrangement of fibers 
to that of Ligniglobus (РІ. 53, figs. 5-8); however, these 
fruits have a scabrate to verrucate surface texture, and 
the pericarp is composed of wider files of cells and has 
abundant large fibrovascular bundles in the pericarp 
(РІ. 59, fig. 7). Woody, unilocular fruits with abundant 
fibrovascular bundles occur in the Palmae. A detailed 
Comparative survey has not been completed. 


Genus SCALARITHECA gen. nov. 


Etymology.— Scalaris (L = of a ladder) + thecium 
= case). 
Туре species.— Scalaritheca biseriata sp. nov. 


Scalaritheca biseriata sp. nov. 
Plate 58, figures 1-9 


Etymology.—Bi (L = two) + series (L = row, suc- 
Cession), referring to the two rows of seeds. 
| Description.—Fruit ellipsoidal, bilaterally symmet- 
Tcal, rounded apically and basally, length 4.7-6.0, avg. 
5.0 mm (SD=0.55, n=7), width 3.0-3.4, avg. 3.1 mm 
(SD=0.2, n=4), thickness 2.8-4.2, avg. 3.6 mm 
(SD=0.57, n=5), fruit bilocular with a thick (0.8-1.0 
Mm) median hollow septum separating two rows of 
elongate seeds oriented with their long axes perpen- 
dicular to the long axis of the fruit; fruit wall smooth, 
0.3-0.4 mm thick, without obvious sclerenchyma; sep- 


2mm 


Text-figure 22. — Scaphicarpium radiatum gen. et sp. nov., fruit in 
transverse section showing mesocarp (grey), endocarp (black), with 
a large placentae and central seed. 


tum rounded ventrally, thickening toward one end; 
seeds elongate, somewhat curved (concave against the 
septum), subcircular to oval in cross section, 2.0-2.4 
mm long, 0.5-0.9 mm wide. 

Specimens.— Holotype: UF 5674. Paratypes: UF 
6107,6281,8779,8780,9603, USNM 355680, 422530, 
435087. 

Discussion.— These silicified structures are inter- 
preted as bilocular fruits with two rows of elongate 
seeds. The familial affinities, however, remain uncer- 
tain. The rows of seeds are visible both in longitudinal 
sections (Pl. 58, figs. 2-4) and in fractured specimens 
(Pl. 58, figs. 1, 6-9). The possibility was considered 
that this species might represent sporocarps of the fern 
Marsilea, with the elongate bodies representing spo- 
rangia, rather than seeds. However, thin sections failed 
to reveal any spores, and the anatomy of the sporangial 
walls in Marsilea, with anticlinally arranged cells, is 
unlike that of the seed coats in this fossil, which have 
periclinally arranged cells. The possibility also exists 
that the fruit-like organ may represent an insect egg 
case, and that the ellipsoidal structures are eggs rather 
than seeds. A throrough comparative check of extant 
insect egg cases was not included in this study. 


Genus SCAPHICARPIUM gen. nov. 


Etymology.— Skaphe (Gr — hollowed out like a boat) 
+ karpos (Gr = fruit), referring to the boat-shaped 
locule casts of this fruit. 

Type species.— Scaphicarpium radiatum sp. nov. 


Scaphicarpium radiatum sp. nov. 
Plate 59, figures 8-17; Text-figure 22 


Etymology.—' The epithet radiatum refers to the 
manner in which fruits radiate from the peduncle. 

Description. —Infructescence a globose head of about 
twenty sessile fruits, head measuring 20 mm in di- 


110 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


ameter; fruit unilocular, single-seeded, ellipsoidal, with 
rounded base and apex, about twice as long as wide, 
circular in transverse section, length 4.7-7.2, avg. 5.5 
mm (SD=0.87, n=8), width 2.6-3.3, аур. 2.7 mm 
(SD=0.80, n=8), thickness 1.9-2.6, avg. 2.2 mm 
(SD=0.28, п-7); pericarp composed of four layers: 1) 
main part of the endocarp wall 200 um thick, com- 
posed of fibers 10-30 um in diameter and more than 
100 um long that are arranged in tracts ranging from 
longitudinal to transverse in orientation; 2) uniseriate 
outer layer of the endocarp, composed of rectangular 
cells 50 um high, 20-30 um wide; 3) mesocarp 300- 
450 um thick, composed of rounded thin-walled iso- 
diametric to somewhat elongate parenchyma cells typ- 
ically 30-60 um in diameter, without ducts or obvious 
vascular tissue; 4) exocarp not preserved. Locule cast 
boat-shaped, rounded-basally, pointed apically, con- 
vex dorsally, concave ventrally, smooth, without a me- 
dian keel, with a prominent circular placental scar cen- 
tral on the ventral face. 


Specimens.—Holotype: USNM 312750 (Bones, 
1979, pl. 6, fig. 5). Paratypes: UF 9523, 9525, USNM 
355368, isolated endocarps; UF 5238 (Bones, 1979, 
pl. 6, fig. 3), 6322, 6324, 6328, 6334, 6359-6363, 
USNM 355458 (2 specimens), 355460 (7 specimens), 
424701, 424731, 446056, 446061, locule casts. 

Discussion. — Scaphicarpium occurs in the Nut Beds 
as permineralized fruits and isolated locule casts. The 
holotype is a silicified infructescence in siltstone matrix 
fractured and showing four attached fruits, plus the 
attachment scars of five more (Pl. 59, fig. 8). Although 
much of the infructescence is hidden with sediment, 
it may be estimated that the full head bore about twenty 
fruits. One fruit from this specimen was removed to 
reveal the locule cast (Pl. 59, figs. 9-12), and was sec- 
tioned transversely (Pl. 59, fig. 13), enabling recogni- 
tion of detached endocarps and locule casts, which are 
relatively common in the Nut Beds. Some of the iso- 
lated specimens proved to be well preserved anatom- 
ically, and revealed anatomy of the endocarp and me- 
socarp (PI. 59, fig. 14-16). 

The boat-shaped locule casts (Pl. 59, figs. 9-13, 17) 
bring to mind Menispermaceae; however, the lack of 
a median keel, and the central, rather than subapical, 
placenta excludes this family. Cantleya in the Icaci- 
naceae also has boat-shaped locules, but the endocarp 
in this genus is two-valved, whereas that of the fossil 
is undivided. I have also considered a possible rela- 
tionship to the Cornales. Fruits in this order are com- 
monly clustered, may be unilocular, and sometimes 
have a boat-shaped locule cast (e.g., Mastixia). How- 
ever, the prominent centrally positioned circular pla- 
cental scar (Pl. 59, fig. 17) and apparent lack of a de- 
hiscence valve is inconsistent with the mastixioids. 


Genus SPHAEROSPERMA gen. nov. 


Etymology.—Sphaero (Gr = ball) + spermum (СТ 
= seed). 
Type species.—Sphaerosperma riesii sp. nov. 


Sphaerosperma riesii sp. nov. 
Plate 58, figures 15-17 


Etymology.—The epithet recognizes the contribu- 
tions of John and Brian Ries to paleobotanical field 
work in the Clarno Formation. 

Description.— Fruit spherical, 24.6-28.5 mm in di- 
ameter, surface smooth, pericarp 1.2-1.4 mm thick 
and composed of: 1) a locule lining about 100 um thick 
of thick-walled rectangular to columnar sclereids, typ- 
ically oriented with their long axes anticlinal and 2) a 
thick layer of rounded, elongate interdigitating cells 
profusely pitted with elliptical pits where they contact 
each other, with intracellular spaces; seed coat thin, 
embryo with two large hemispherical cotyledons. 

Specimens.—Holotype: UF 9746. Paratype: ШЕ 
9599. 

Discussion. —Sphaerosperma is represented by two 
specimens, both of which are embedded in hard tuff- 
aceous matrix so that the outer shape is apparent only 
by fractures and sections (Pl. 58, figs. 15, 16). The 
holotype appears to show two hemispherical cotyle- 
dons. It is similar in shape and size to Coryloides, but 
has a smooth, rather than ribbed outer surface, lacks 
the cupule scar, and has a wall that is anatomically 
preserved, evidently more resistant to decay than that 
of Coryloides. The interdigitating cells composing the 
endocarp wall resemble those of Palaeophytocrene (cf. 
РІ. 58, fig. 17 and Pl. 20, fig. 5), but whether this is an 
indication of Icacinaceous affinity, or simply a case of 
convergence, or has not been determined. 


Genus SPHENOSPERMA gen. nov. 


Etymology.— Spheno (Gr = wedge) + spermum (Gr 
= seed). 
Type species.— Sphenosperma baccatum sp. nov. 


Sphenosperma baccatum sp. nov. 
Plate 60, figures 7-10; Text-figure 23 


Etymology — The epithet is derived from bacca (L ^ 
berry). | 

Description.—Fruit subglobose, prolate, circular 10 
cross section, length 13 mm (est.), equatorial diameter 
9.2 mm, surface smooth, unilocular with three partial 
septa 200-500 и thick extending from the periphery of 
the fruit almost to the central axis, without an obvious 
central column, and with six radially arranged wedge 
shaped seeds (five fully developed, one small); репсатр 
200-300 и thick, of two layers: 1) the inner Уз to № 
with horizontally oriented fibers 15-25 um wide, 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 111 


the % to % with anticlinally oriented fibers 12-15 um 
Wide. Seeds nearly triangular in cross section, with a 
Сопуех dorsal surface and straight ventral sides that 
Meet at 65-85°, with a “swelling” along Ше ventral 
Side containing the raphe, seed height similar to that 
of fruit, width 3.3—4.2 mm, dorsiventral thickness 3.3— 
4.2 mm; seed coat 200 um thick consisting of two 
layers: 1) the inner layer five- to seven-seriate, com- 
Posed of rounded more or less isodiametric cells 20- 
50 um in diameter, 2) the outer layer uniseriate, com- 
Posed of large, columnar, anticlinally oriented cells 80- 
1 20 um high, 25-45 um wide with concave outer walls, 
8lving the outer surface of the seeds a scalloped ap- 
Pearance. 

Specimen.—Holotype: UF 5666. 

Discussion.— Sphenosperma is known from a single 
зресипеп that was exposed in fracture planes through 
Indurated siltstone, so that Ше exterior surface could 
Not be photographed directly. The pieces were glued 
together so that the fruit could be sectioned transverse- 
ly (Pl. 60, figs. 7-10). Successive serial sections re- 
vealed the morphology described above. 

Important features of this taxon include the three 
Septa (suggesting a tricarpellate fruit) and the wedge- 
Shaped seeds with the punctate surfaces due to col- 
lapsed external walls of the outer layer of seed coat. I 
Suspect that the number of seeds per fruit was variable 
Ш this species. In this specimen there are three seeds 
(including one that is abortive) between two adjacent 
Septa, but only two and one seed are situated, respec- 
Uvely, between the other two pairs of adjacent septa. 


Genus STOCKEYCARPA gen. nov. 


Etymology.—This genus is named for Ruth A. 
tockey, recognizing her contributions to paleobotany. 
Type species.— Stockeycarpa globosa sp. nov. 


Stockeycarpa globosa sp. nov. 
Plate 60, figures 11—14 


| Etymology.— Globus (L = ball), referring to the fruit 
аре. 

Description. —Fruit spherical, unilocular, base and 
арех rounded, 6.8 mm in diameter, smooth-walled; 
Inner layer composed of anticlinally elongate columnar 
Cells 40-45 um high, 6-8 um wide, middle layer 400— 
200 um thick composed of parenchyma apparently with 
tracellular spaces, outer preserved layer 300 um thick, 
"hiseriate, composed of anticlinally oriented fibers 300 
Hm high, 25-38 um wide. 

Specimen.— Holotype: UF 9747. 

Discussion. — Stockeycarpa is represented by a single 
SPecimen in the Nut Beds. Although the seed is not 
Preserved, the fruit anatomy is distinctive with the two 
ayers of anticlinally elongate cells separated by a pa- 


Text-figure 23.—Sphenosperma baccatum gen. et sp. nov., trans- 
verse section of the fruit showing six seeds (one small, perhaps abor- 
tive), and three intruding partial septae. 


renchymatous mesocarp (PI. 60, figs. 13, 14). Another 
specimen that appears to be virtually identical mor- 
phologically and anatomically has been recovered in 
the Eocene Princeton Chert of British Columbia (Cev- 
allos-Ferriz, pers. comm., 1992). 


Genus STRIATISPERMA gen. nov. 


Etymology. — Stria (L = line, furrow) + sperma (Gr 
= seed). 

Type species.— Striatisperma coronapunctatum sp. 
nov. 


Striatisperma coronapunctatum sp. nov. 
Plate 60, figures 1-6 


Etymology.— Corona (L = crown) + punctum (L = 
small hole, dot). 

Description.—Seed globose to ellipsoidal, some- 
times unevenly contorted or compressed, typically cir- 
cular in cross section, rounded apically and basally, 
length 4.1-9.6 (-14.0), avg. 6.1 mm (SD=1.45, п=59), 
equatorial diameter 2.7-8.5 (-11.2), avg. 5.7 mm 
(SD=1.20, n=58), with a circular scar at base and a 
small projection at the apex encircled by a cycle of 
about ten small circular to elliptical protrusions or de- 
pressions 60—100 um in diameter, surface prominently 
striate with longitudinal rows of rectangular cells 8—11 
шт in diameter, outer striate layer sometimes partially 
broken away and revealing another similarly striate 
surface with the same cell pattern. Sometimes with 
funicle or placental strands linking two or more seeds. 

Specimens.— Holotype: USNM 424694. Paratypes: 
OMSI Pb271, UCMP 10624, 10625, UF 6325, 6335, 
6455, 8700-8727, 8809-8815, 9211, 9212, USNM 
355053 (3 specimens), 355054 (2 specimens), 355064- 


112 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


355068, 355069 (4 specimens), 355070 (5 specimens), 
355071, 355633, 422385, 424695, 424696, 424873, 
435055, 435056. 

Discussion. — Striatisperma seeds are relatively com- 
mon at the Nut Beds. They are distinctive because of 
the outer surface oflongitudinally aligned rows ofsquare 
to rectangular cells (Pl. 60, figs. 1-6), and the ring of 
circular protrusions or scars surrounding the apex. The 
function of the circular scars is not known. Although 
typically more or less ellipsoidal and about 6 mm long, 
the seeds are variable in shape and size, some speci- 
mens being more than a centimeter in length. One 
specimen, UF 6455, is particularly large, 14.0 mm 
long, 10.0 x 11.2 mm wide. Although some examples 
have been sectioned and some are broken, no internal 
tissue has been observed beneath the two outer layers 
of longitudinally aligned cells. One specimen shows 
two silica seed casts partially embedded in matrix, 
linked by the silica cast of a bent, cylindrical funicle 
or placental strand (Pl. 60, fig. 5). 


Genus TENUISPERMA gen. nov. 


Etymology.— Tenuis (L = thin) + spermum (Gr = 
seed), referring to the thinness of the seed. 
Type species.— Tenuisperma ellipticum sp. nov. 


Tenuisperma ellipticum sp. nov. 
Plate 61, figures 1-9 


Etymology.— Ellipsis (L = a defective circle; ellipse) 
referring to the seed outline. 

Description.—Seed elliptical in face view, dorsiven- 
trally flattened and somewhat sinuous or wavy, base 
rounded, apex pointed; length 6.3, 10.5 mm, width 
5.5, 7.2 mm, thickness 2.0, 2.0 mm, seed coat smooth, 
250 um thick, with an inner uniseriate layer of anti- 
clinally elongate columnar cells 120-140 um high, 30- 
60 um wide and an outer three- to four-seriate layer 
composed of isodiametric sclereids, 30-40 um in di- 
ameter; lateral margin with a grooved ridge extending 
from a point 2 mm from the apex on one side around 
the base and up to the position near the apex on the 
opposite margin of the seed. 

Specimens.—Holotype: UF 6108. Paratype: UF 
9528. 

Discussion.— Tenuisperma seeds are preserved as 
white chalcedony seed casts. A ridge on the lateral 
margin has a median groove passing from near the 
apex on one side, over the base and to near the apex 
on the opposite margin of the seed. This groove ter- 
minates just prior to the top of the ridge, giving the 
appearance of a doubled, U-shaped structure (Pl. 61, 
fig. 4). This structure may represent the edges of the 
two cotyledons that are planar and closely adpressed 
to one another. Thus the seed is exalbuminous, filled 
by the embryo. i 


Genus TIFFNEYCARPA gen. nov. 


Etymology.— Tiffney + karpos (Gr = fruit). The ge- 
nus is named in honor of Bruce H. Tiffney, paleocar- 
pologist. 

Type species.— Tiffneycarpa scleroidea sp. nov. 


Tiffneycarpa scleroidea sp. nov. 
Plate 62, figures 1-7 


Etymology.— Skleros (Gr = hard, tough), referring 
to the woodiness of the fruit. 

Description. — Fruit subglobose, prolate, loculicidal 
capsule; base bluntly rounded with a central depression 
0.95 mm in diameter; apex unknown, length 13 mm, 
equatorial diameter 9.5 mm; surface with longitudinal 
ribs and grooves, fruit woody, thick-walled with ten 
radially fused carpels and eight developed locules, each 
with a single seed; central axis with about ten vascular 
bundles alternating with the locules; locules vertically 
elongate, up to ca. 8 mm high, 0.8-1.3 mm wide, 1.7- 
2.3 mm dorsiventral dimension, locule lining unise- 
riate, composed of fibers 15—20 um in diameter; en- 
docarp composed mostly of equiaxial sclereids with 
undulating margins, 50-80 um in diameter; radial 
planes of dehiscence formed by smaller sclereids ra- 
diating from the central axis and bisecting each locular 
area; exocarp (or possibly mesocarp) forming a band 
400—500 um thick of anticlinally oriented polygonal 
cells of variable size 40-120 um radial dimension, 30- 
80 um wide. Seeds attached to the ventral margin О 
the locule near the apex, embryo straight. 

Specimen. — Holotype: USNM 355645. 

Discussion. — Tiffneycarpa is known only from а sin- 
gle specimen that shows evidence of predepositional 
waterwear (Pl. 62, figs. 1-3). It is partially broken and 
replaced with silicified sandstone. Although the арех 
is missing, anatomical preservation is excellent, 45 seen 
in serial transverse thin sections (Pl. 62, figs. 4—7). 

Distinctive features of this fruit are the woodines5, 
ten carpels, subapical axile placentation, and loculi- 
cidal planes of dehiscence (or carpel fusion) that extend 
fully from the axis to the endocarp periphery (Pl. 62, 
figs. 5-7). The morphology and anatomy are reminiS” 
cent of the extinct genera Wetherelia and Palaeoweth- 
erelia (see Mazer and Tiffney, 1982). However, these 
genera have both loculicidal and septicidal dehiscence 


Genus TRIGONOSTELA gen. nov. 


Etymology.—Trigono (Gr = triangular) + stele (Gt 
= column). 
Туре species.— Trigonostela oregonensis sp. NOV- 


Trigonostela oregonensis sp. nov. 
Plate 61, figures 10-18 


Etymology.—The epithet refers to Oregon. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 113 


Description.—Fruit elongate, tricarpellate, trisym- 
metrical with three planes of symmetry intersecting in 
the long axis, lateral walls disposed in a trigonal pris- 
Matic pattern, trigonal in cross section, oblong in lon- 
gitudinal view, apex pointed, base blunt, length of 
Complete fruit 5.5-6.1, avg. 5.8 mm (SD=0.28, n=4), 
Width 2.6-2.9, avg. 2.7 mm (SD=0.13, n=4); length 
Of locule cast 4.8-5.4, avg. 5.2 mm (SD=0.32, n=3), 
Width 1.7-2.0, avg. 1.9 mm (SD=0.17, n=3); three 
lateral faces joining to form longitudinal ribs, each of 
the three faces with a median groove corresponding to 
the position of a septum. 

Specimens.—Holotype: USNM 424671. Paratypes: 
UF 6297, USNM 354599, 355643, 424672-424675, 
435042. 

Discussion.— Trigonostela is represented by silica 
Casts and partial permineralizations. The elongate tri- 
80nal form of this fruit (Pl. 61, figs. 10-16) is very 
distinctive; however, the affinities have not been de- 
termined. One of the specimens sectioned transversely 
Shows three septa that intrude inward, but which do 
Not intersect, indicating a unilocular fruit (Pl. 61, fig. 
17). Another specimen that was sectioned transversely 
appears to be composed of radiating fibers with poorly 
developed locules (Pl. 61, fig. 18). The anatomy and 
Number of seeds per locule are not known. 


Genus TRIPARTISEMEN gen. nov. 
Etymology.— Tripart (three-parted) + semen (Г. = 
Seed). 
Type species.— Tripartisemen bonesii sp. nov. 


Tripartisemen bonesii sp. nov. 
Plate 62, figures 8-15 


Etymology.— This species is named after Thomas J. 
Ones, who collected most of the specimens. 

| Description.—Seed subglobose, bilaterally symmet- 
пса], dorsal surface rounded, ventral side with two 
acets joining at ca. 110? to form a median longitudinal 
Crest in the plane of symmetry; with a pair of longi- 
tudinal keels formed at the junction of the ventral fac- 
“ts with the dorsal surface (In cross section subtrian- 
Sular with a rounded dorsal side and a pair of straight 
Ventral sides); base rounded, with a small circular hilar 
Scar, apex pointed; seed length 2.8-4.4, avg. 3.5 mm 
(SD=0.41,n=31), width across plane of symmetry 2.6— 
4.3, avg. 3.4 mm (SD=0.43, n=31), thickness 2.1-3.5, 
аур. 2.8 mm (SD=0.35, n=20); seed coat smooth, sep- 
tum 100-200 um thick, Y-shaped, dividing the seed 
Into three chambers: a pair of large, apparently empty, 
ateral chambers and a slender median, dorsal embryo 
Chamber: embryo chamber sickle-shaped, curved in 
Ше plane of symmetry, with a flattened, rounded-tri- 
angular facet at the base, possibly the chalaza or a 
Sermination valve; septum formed of small isodia- 


metric cells 12-20 um in diameter, with an outer layer 
of cuboidal to rhomboidal thin walled cells 8-10 um 
thick. 

Specimens.— Holotype: UF 9288. Paratypes: UCMP 
10741-10742, UF 9289-9316, 9577, 9578, 9734 (100 
in lot), USNM 354484 (4 specimens), 354485 (18 spec- 
imens), 354486 (84 specimens), 354487 (4 specimens), 
354488, 355688 (in matrix), 355395 (14 specimens), 
355445 (5 specimens), 355688 in matrix. 

Discussion.— Tripartisemen bonesii is one of the most 
abundant species in the Nut Beds, yet it is not known 
from other localities of the Clarno Formation or else- 
where in North America. It is often preserved as casts 
of the three chambers which may easily fall apart when 
the specimen is exposed. Permineralized seeds have 
been sectioned to reveal anatomy of the walls and septa 
(Pl. 62, figs. 14, 15). Sometimes the lateral chambers 
are filled with sediment, suggesting that they were orig- 
inally empty, but had ruptured. The central chamber 
is generally filled with translucent silica, not clastic 
sediment, and in at least one instance (Pl. 62, fig. 15), 
preserves an embryo. 

Collinson (1988) illustrated this type of seed both 
from the Nut Beds and from the Eocene of Messel, 
Germany. Although the well-preserved, lignitized 
Messel specimens differ in the nature of their preser- 
vation from the silicified Clarno seeds, the morphology 
appears to be identical. Collinson (1988 and work in 
progress) suggests possible lythraceous affinities for this 
genus. Tripartisemen resembles the extinct lythraceous 
genus Mneme Eyde in symmetry and presence of a 
median and two lateral chambers. However, there are 
a number of differences, including that Mneme seeds 
are about half as large, the embryo chamber is larger 
than the lateral chambers, and the lateral chambers are 
relatively thick-walled and are open along median slits 
(Friis, 1985). 


Genus TRIPLASCAPHA gen. nov. 


Etymology.— Tripla (L = triple) + scapha (L = small 
boat), referring to the three boat-shaped locules of the 
fruit. 

Type species.— Triplascapha collinsonae sp. nov. 


Triplascapha collinsonae sp. nov. 
Plate 63, figures 1-12; Text-figure 24 


Etymology.— The epithet recognizes the important 
contributions of Margaret E. Collinson to paleocar- 
pology. 

Description.— Fruit ovoid, tricarpellate, trisym- 
metrical, nearly circular in cross section, one end (base?) 
rounded, the other (apex?) truncate with a circular disk; 
fruit length 4.0-6.0 mm, equatorial diameter 3.7-5.5 
mm, trilocular with a single seed per locule; locule casts 
shallowly c-shaped in cross section, the ventral face 


114 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


5mm 


Text-figure 24. — Triplascapha collinsonae gen. et sp. nov., trans- 
verse section showing three seeds within parenchymatous pericarp. 


convex, the dorsal face concave, 4.1 mm long, 3.0-3.1 
mm wide. Pericarp consisting of four layers: 1) locule 
lining five- to 12-seriate, composed of isodiametric to 
periclinally and horizontally elongate cells 10-15 um 
thick and up to 33 um long, 2) a uniseriate layer of 
columnar cells 55 to 140, avg. 80 um high and 15-23 
um wide, 3) a 50 um-thick layer of horizontally ori- 
ented narrow fibers each about 2.5-3.8 um thick; layers 
1-3 defining three pyrenes, which together are sur- 
rounded by 4) a layer of parenchyma cells 30-58 um 
in diameter, 0.7-2.0 mm thick. Embryo circular in 
cross section, tube-like, aligned with the center of each 
locule. 

Specimens.—Holotype: UF 9472. Paratypes: UF 
9480, USNM 354051. 

Discussion.—Triplascapha is represented by rare 
locule casts and permineralized fruits. The fruits are 
distinctive in the presence of a circular disk-like trun- 
cation at one end, possibly the apex (Pl. 63, figs. 1, 2, 
7, 8), and the three dorsally concave locules that are 
symmetrically arranged about the long axis of the fruit 
(Pl. 63, fig. 10; Text-fig. 24). Typically, the outer layers 
of the fruit are stripped away as they are broken from 
the sediment, but one specimen still attached to the 
sedimentary matrix was sectioned to show the full 
thickness of mesocarp (Pl. 63, fig. 9). In size and form 
of the endocarp and locules, this fruit converges closely 
with Portnallia Reid et Chandler, a genus from the 
London Clay that has been synomized within the mas- 
tixioid genus Beckettia (Knobloch and Mai, 1986). For 
a detailed anatomical comparison, I was permitted to 
cut a transverse section of Portnallia bognorensis 
(v.30422, a specimen originally figured by Reid and 
Chandler, 1933, pl. 26, fig. 41). This section reveals 
that the London Clay endocarp is composed of swirling 
tracts of fibers, and lacks a central vascular bundle— 


characters that corroborate its assignment to the mas- 
tixioids and differentiate it from Triplascapha. 

Transverse sections of Triplascapha reveal a dis- 
tinctive columnar layer surrounding each of the three 
locules (Pl. 63, figs. 11, 12) that is not seen in extant 
or fossil mastixoids. The fibrous layer surrounding the 
columnar cell layer around each locule appears to de- 
fine a pyrene, so that the three endocarps might have 
been shed independently unlike the situation in Cor- 
naceae. The cylindrical, tube-like embryo (Pl. 63, figs. 
10, 11) is a peculiarity that may eventually help in 
pinning down affinities with a particular modern fam- 
ily. 

Genus TRIPLEXIVALVA gen. nov. 


Etymology.—Triplex (L = threefold) + valva (L = 
door), referring to the three valves of the fruit. 
Type species.— Triplexivalva rugata sp. nov. 


Triplexivalva rugata sp. nov. 
Plate 63, figures 13-20 


Etymology.— Rugosus (L = wrinkled). 

Description.— Fruit subglobose, nearly circular tO 
rounded-triangular in cross section, rounded basally 
and apically, with a circular scar at the base, and à 
triangular to circular scar at apex; length 3.3-8.1, ауё. 
5.3 mm (SD- 1.37, n=11), width 3.5-7.7, avg. 5.3 mm 
(SD=1.44, n=10), narrower width 2.8-7.5, avg. 4.8 
mm (SD- 1.83, n=6); composed of three well-defined 
valves, whose margins meet to form three longitudinal 
ridges that intersect apically and basally; asymmetrical, 
often with one valve somewhat larger than the others, 
valves up to 0.8 mm thick; surface longitudinally ru- 
gulate, rugulae thinning apically. Locule casts trigonal 
in transverse view, with the three carpels radially fused, 
sometimes with a cleavage between them, otherwise 
with a flange indicating the edges of adjoining valves- 

Specimens.— Holotype: USNM 424799. Paratypes: 
UF 5237 (Bones, 1979, pl. 5, fig. 9), 6378—6380, 6382. 
9725-9727, USNM 312759 (Bones, 1979, pl. 5, fig. 4). 
355539 (2 specimens), 422396, 424798. 

Discussion.— This species is represented by trigonal, 
globose fruit casts with prominent longitudinal ribs (Pl. 
63, figs. 13-20). Circular spots on the surface of some 
specimens (Pl. 63, figs. 13, 15) might represent an ех- 
udate. Some of Ше specimens have shed one or more 
valves, showing that the valves themselves are rela- 
tively thick, and revealing a smooth locule cast inside 
(Pl. 63, figs. 17-20). 


TRISEPTICARPIUM gen. nov. 


Etymology.—Tri (L, Gr = three) + septum (L = 
partition) + karpos (Gr = fruit). 
Type species.— Trisepticarpium minutum sp. поу. 


| 
| 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER VAS 


Trisepticarpium minutum sp. nov. 
Plate 64, figures 1-10 


Etymology.—The epithet refers to the small size of 
the fruit. 

Description.—Fruit pyriform, circular in cross sec- 
tion, base cuneate, rounded, apex hemispherical, 
Tounded; length 2.8 mm, width 2.2 mm, tricarpellate, 
trilocular; surface smooth or faintly striate longitudi- 
nally, with three longitudinal grooves intersecting at 
the apex; outer wall of fruit 140 um thick, composed 
of densely packed more or less isodiametric sclereids 
25-50 um in diameter, ventral wall of each carpel 
thicker than the outer fruit wall and comprised of a 
Uni- to biseriate layer of large anticlinally elongate co- 
lumnar cells 180-400 um high, 40-100 um wide; inner, 
ventral, wall of each carpel joining with those of ad- 
Jacent carpels to form a prominent triradiate to tri- 
angular septum, locule lining composed of a uniseriate 
layer of horizontally oriented fibers 50 um thick; loc- 
ules dorsiventrally compressed 259-300 um thick and 
1150-1250 um wide, containing small winged seeds, 
Placentation axile and near the apex, seeds in cross 
section with a lensoidal central body 220-280 um wide, 
80 um thick covered by a finely reticulate layer which 
is drawn out into a thin wing on both sides, width 
including wings 450 um, wing width approximately as 
long as the width of the central body, straight or re- 
Curved near the margins, seeds arranged in two rows 
рег locule, apparently with central placentation. 

Specimen.— Holotype: USNM 424737. 

Discussion.—This new genus and species is repre- 
Sented by a single well-preserved specimen which was 
Photographed in stereo (Pl. 64, figs. 2, 3) and then 
Sectioned transversely to reveal well preserved anat- 
Omy. The thick inner septa of this fruit, joined in a Y 
Configuration (Pl. 64, figs. 5-9), and the winged seeds 
(Pl. 64, fig. 10), are distinctive features of this fruit. 


Genus TRUNCATISEMEN gen. nov. 


Etymology.— Truncus (L = cut off) + semen (L = 
Seed). 


Type species.— Truncatisemen sapotoides sp. nov. 


Truncatisemen sapotoides sp. nov. 
Plate 64, figures 11-14 


Etymology.—The epithet refers to the similarity with 
Seeds of Sapotaceae. 

Description.—Seed ellipsoidal, bilaterally symmet- 
rical, laterally compressed in the plane of symmetry 
With an oblique apical truncation, base and lateral mar- 
gins smooth and rounded, length 9.5-12.2 mm, width 
In plane of symmetry 6.2-8.0 mm, thickness across 
Plane of symmetry 5.5 mm; surface smooth, shiny, 


truncate face elliptical, 2.7-3.0 mm wide, 5.2—5.8 mm 
long, forming a slope about 35—40° to the long axis of 
seed, with a circular scar (hilum) in the center. 

Specimens.— Holotype: UF 6459. Paratype: USNM 
424650. 

Discussion.— Both specimens of this species are pre- 
served as chalcedony casts. Cracks on the surface of 
both fossil seeds suggest a thin, easily fractured seed 
coat. Similar obliquely truncate seeds occur in Styra- 
caceae, Illiciaceae and Sapotaceae, however the precise 
familial affinities remain undetermined. Although there 
is a gross resemblance to seeds of some Sapotaceae, 
Truncatisemen does not show the characteristic bulg- 
ing scar beneath the truncation that is typical of Sa- 
potaceae. Т. sapotoides differs from the Clarno truncate 
seeds attributed to Bumelia? by larger size, and lateral 
compression. 


Genus ULOSPERMUM gen. nov. 


Etymology.— Ulo (Gr = scar) + spermum (Gr = seed), 
referring to the prominent attachment scar. 
Type species. — Ulospermum hardingae sp. nov. 


Ulospermum hardingae sp. nov. 
Plate 65, figures 1-8 


Etymology.— This species is named after Elisabeth 
Harding, who collected one of the best specimens. 
Description.—Fruit bilaterally symmetrical, 
obliquely cordiform in the plane of symmetry, with a 
large concave, elliptical scar at the broad end, pointed 
at the opposite end, oblong in cross section; fruit length 
13.2-14.3, avg. 13.9 mm (SD=0.61, n=3), width 16.4- 
18.0, avg. 17.0 mm (SD=0.87, n=3), thickness across 
plane of symmetry 8.8-11.0, avg. 9.6 mm (SD=1.17, 
n=3), length/width ratio 0.73-0.87, avg. 0.82; the el- 
liptical scar 6 х 9 mm, containing many (about 20) 
protruding V- or Y-shaped vascular traces; wall 0.8 
mm thick, made up of three anatomical layers: 1) uni- 
to biseriate locule lining ca. 30 um thick composed of 
thin walled poorly preserved periclinally elongate rect- 
angular cells 12-15 um in diameter, 2) a middle layer 
ca. 250 um thick composed of anticlinally organized 
files of densely packed small sclereids 12-22 um wide, 
30—38 um long, grading into, 3) outer layer composed 
of larger cells 40-80 (-130) um in diameter that are 
not anticlinally aligned, grading from more or less iso- 
diametric to larger and periclinally elongate toward the 
periphery. Locule cast relatively smooth, in one spec- 
imen bearing small, evenly spaced punctae. 
Specimens.— Holotype: USNM 355363. Paratypes: 
UF 9566, USNM 414509, 355363. 
Discussion.— This species is rare in the Nut Beds 
collections and is unknown from other localities. Two 
of the specimens are locule casts with the outer wall 


116 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


missing (e.g., Pl. 65, figs. 1-3). The tissue over the large 
recessed scar appears to have been well-indurated, and 
has remained intact in one specimen (Pl. 65, fig. 1), 
but when removed, shows numerous vascular bundle 
endings (Pl. 65, figs. 2-4). The holotype has the wall 
intact, permineralized, and, upon sectioning, yielded 
the anatomical details described above (Pl. 65, figs. 5- 
8). Although Ulospermum could be interpreted either 
as a seed with a conspicuous hilum, or as a fruit with 
a conspicuous cupule or pedicel scar, I have described 
it as a fruit, because the specimen that was cut trans- 
versely shows the remains of what appears to be a seed 
shriveled in the center of the locule (Pl. 65, fig. 5). 
Despite the very distinctive morphology, I have been 
unable to determine its affinity among extant families. 


Genus WHEELERA gen. nov. 


Etymology.— This genus is named for Elisabeth A. 
Wheeler, recognizing her contributions to paleobotany 
including research on woods of the Clarno flora. 

Type species.— Wheelera lignicrusta sp. nov. 


Wheelera lignicrusta sp. nov. 
Plate 65, figures 9-16 


Etymology.— Lignum (L = wood) + crusta (L = 
shell), referring to the fibrous fruit wall. 

Description.— Fruit subglobose, prolate, rounded 
basally and apically, unilocular, length 11.4-12.0 mm, 
equatorial diameter 8.0-8.4 mm; pericarp 1.4-2.0 mm 
thick, composed of: 1) uniseriate locule lining of thick- 
walled anticlinally oriented columnar cells 100 um high, 
25 um wide that commonly contain one to four rhom- 
boidal crystals, 2) a thick layer of short fibers 300-600 
um long, 40—70 um wide, 3) an outer layer about 200 
um thick composed of tangentially flattened of paren- 
chyma cells 60-100 um wide, 13-40 um high. Locule 
cast smooth, rounded basally, pointed apically. 

Specimens.—Holotype: UF 9271. Paratypes: UF 
9576, 9359. 

Discussion. — This species is known from three per- 
mineralized specimens. The main thickness ofthe peri- 
carp formed of anticlinally directed short fibers (Pl. 65, 
fig. 15), and the locule lining of columnar cells with 
crystals (Pl. 65, fig. 16) are distinctive features of this 
genus. As yet, the affinities with modern taxa remain 
undetermined. 


Genus CARPOLITHUS Schlotheim 


Carpolithus bellispermus Chandler 
Plate 66, figures 1-10 


Carpolithus bellispermus Chandler 1978, pp. 31-32, pl. 8, figs. 3-6. 


Description.—Seed ovate in face view, dorsiven- 
trally compressed, rounded-triangular in cross section, 


typically bilaterally symmetrical except at the laterally 
oblique apex; base rounded, apex in complete seeds 
rounded, slightly bulging, with an asymmetrical, an- 
gular notch visible on the dorsal and ventral surfaces; 
dorsal side of seed gently convex, ventral side typically 
with two faces meeting at about 120° to form a rounded 
median longitudinal ridge; length of complete seed 2.8- 
3.2, avg. 3.0 mm (SD=0.19, n=7), width 1.5-2.0, avg. 
1.8 mm (SD=0.17, n=7), thickness 0.9-1.1, avg. 1.0, 
(SD=0.08, n=7); seed coat composed of two cell layers: 
inner layer uniseriate, about 0.02 mm thick, composed 
of small, rectangular to polygonal cells more or less 
longitudinally aligned as viewed peridermally; outer 
layer 0.125 mm thick, uniseriate with large cells that 
are adjoining basally, but with each cell narrowed api- 
cally and forming a cylindrical digitate projection, dig- 
itations about 30 um in diameter, oriented anticlinally, 
resulting in a papillate surface to the seed coat; digi- 
tations are encased in a tan to brownish translucent 
substance, possibly wax, that forms the outer surface 
of the seed. Endosperm cast (internal mold of seed 
coat) shaped like the complete seeds, except obliquely 
truncate apically, the plane of truncation oriented 45- 
60° to the longitudinal axis as viewed ventrally or dor- 
sally; micropyle forming a prominent point at the top 
of the apical truncation. 

Specimens.—UCMP 10663-10667, 10698-10700; 
USNM 355416, 435014-435017; UF 6503-6506, 
8544-8579, casts; UCMP 10696, 10697, UF 6607, 
specimens with intact seed coat. Holotype: BM v5 1663 
(London Clay). 

Discussion.— Carpolithus bellispermus was recog- 
nized previously only from the Eocene Bognor Beetle 
Bed of the London Clay, England (Chandler, 1978). 
The familial affinities are still unknown, and the species 
is retained provisionally in the noncommittal genus 
Carpolithus. It is represented in the Nut Beds by nu- 
merous internal chalcedony molds of seed coat, and à 
few permineralized specimens with the seed coat in- 
tact. The internal molds are readily recognized by the 
smaller size, obliquely truncate apex, rounded Базе, 
dorsiventral compression and median ventralridge (Pl. 
66, figs. 5, 6). Complete seeds show the notched apex 
(Pl. 66, figs. 1-4) and digitate outer layer (Pl. 66, figs. 
7-9). The closely spaced, anticlinally oriented digitate 
cells, enveloped in a translucent layer, are conspicuous 
features that should be helpful in the search for related 
extant and/or fossil taxa. Despite good preservation of 
the seed coat in some of the specimens, no embryo 
and no raphe were observed. 

In the description above I have followed Chandler’s 
(1978) interpretation that these specimens represent 
seeds rather than fruits. Chandler (1978) provided 4 
brief description of fruit characters based upon зрес- 


| 
| 


| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 117 


imen(s) that she had observed prior to pyritic deteri- 
Oration. She stated that only part of the endocarp was 
Preserved, bearing a single seed and having a longi- 
tudinal suture with rounded edges along which splitting 
had occurred and noted that the was wall columnar in 
Section. She interpreted the seeds to be campylotro- 
Pous because of the adjacent hilum and micropyle 
without an obvious raphe. Corresponding fruits have 
not been recovered from the Nut Beds, but the seeds 
are so distinctive that there can be no question in iden- 
tifying this material to the London Clay species. 


Carpolithus sp. 1 
Plate 66, figures 11-14 


Description.—Endocarp unilocular, ellipsoidal, 
nearly circular in cross section, 7.8-11.0 mm long, 6.1— 
6.9 mm in diameter, base and apex rounded, surface 
longitudinally ribbed, with about eight to ten promi- 
nent, rounded meridional ridges, each with a groove 
at its crest; endocarp wall 0.7 mm thick. 

Specimens. — USNM 355686, ОМІ Pb1439. 

Discussion.— This eight- to ten-ribbed fruit is rep- 
Tesented by two specimens from the lower part of Face 
3. Neither is anatomically preserved, but the external 
Morphology indicates that it represents a species dis- 
tinct from any other known from the Nut Beds. 


Carpolithus sp. 2 
Plate 66, figures 15, 16 


Description.— Achene bisymmetrical, ovate in lat- 
tral view, base truncate, apex acute; laterally com- 
Pressed in the plane of symmetry with a wing-like keel 
On one side, rounded on the other side; length 2.4–2.7 
mm, width 1.6-1.7 mm, thickness 1.0-1.1 mm; trun- 
Cate base with an elliptical recessed area (hilum or 
pedicel scar), surface smooth on the rounded margin, 
but cratered with shallow depressions up to 200 um in 
diameter on the lateral margins and on the wing-like 
Portion. 

Specimens.—USNM 424719, UF 8581. 

Discussion.—This species is represented by hollow 
Silica casts that appear to represent achenes. Distinc- 
tive features are the relatively large circular basal scar 
(PL 66, fig. 16), the cratered surface and wing-like keel 
On one side. 


Carpolithus sp. 3 
Plate 66, figures 17, 18 


Description.—Seed prolate, ellipsoidal, circular-el- 
liptical in cross section, base and apex rounded, ellip- 
tical in cross section, length 6.4 mm, diameter at equa- 
tor 5.0 x 5.8 mm, surface smooth; seed coat 280 um 
thick, composed of: 1) an inner layer ca. 200 um thick 
Of more or less isodiametric parenchyma cells 13-30 


um in diameter, 2) a middle layer ca. 50 um thick of 
tangentially flattened sclerenchyma, 3) an outer layer 
about 40 um thick composed of larger thin-walled an- 
gular parenchyma cells 30-50 um in diameter. Coty- 
ledons large, plano-convex, hemispherical. 

Specimen.—USNM 355069. 

Discussion.—This species is represented by a single 
specimen. Although the outer surface is rather non- 
descript, sectioning revealed a thick wall and large pla- 
noconvex cotyledons (Pl. 66, fig. 18). Although similar 
to Lauraceae in the single locule and large hemispher- 
ical cotyledons, the columnar layers typical of that fam- 
ily were not observed. The preserved wall is interpreted 
above as representing the seed coat, but it may also 
include part of the fruit wall. 


Carpolithus sp. 4 
Plate 67, figures 1-5 


Description.—Locule cast obpyriform, bisymmetri- 
cal, nearly circular in cross section, but with a raised 
longitudinal ridge in the plane of symmetry, base and 
apex pointed, widest about У of the distance below 
apex, length 5.0 mm, diameter 2.0 mm; the longitu- 
dinal ridge formed only on one side in the plane of 
symmetry, appearing as a flattened band 0.7 mm wide 
with a median keel; single-seeded with thin seed coat 
preserved. 

Specimen.—USNM 424652. 

Discussion.— This specimen appears to be the locule 
cast ofa unilocular fruit. The longitudinal ridge, which 
is only on one side (ventral?) apparently represents the 
suture plane (Pl. 67, figs. 1-4). A transverse section 
reveals a single seed inside with a thin seed coat pre- 
served (Pl. 67, fig. 5). At least superficially, it resembles 
the locule of Prunus (Mai, pers. comm., 1989), but it 
does not correspond to the locule morphology of either 
of the Prunus species known from permineralized en- 
docarps from the Nut Beds. 


Carpolithus sp. 5 
Plate 67, figures 6-8 


Description.—Fruit unilocular, single-seeded, en- 
docarp more or less ellipsoidal, elliptical in cross sec- 
tion, length 3.4 mm, width 3.1 mm, thickness 2.0 mm; 
endocarp wall smooth, 230—330 um thick, ca. 12-se- 
riate, composed of rounded cells with small extensions, 
adjacent cells adjoining like pieces of a jig-saw puzzle, 
these cells typically 15-40 um in diameter; exocarp 
uniseriate with inflated, thin-walled rectangular cells, 
25-58 um wide, 20-38 um high, and an outer biseriate 
layer of square to rectangular cells 10-13 um in di- 
ameter. Seed coat uniseriate, composed of cuboidal 
cells about 10 um in diameter. 

Specimen.— UF 9758. 


118 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Discussion. — This species is represented by a single 
well-preserved specimen. It was sectioned transversely 
revealing the pericarp thickness and a single seed with 
a shriveled thin seed coat (Pl. 67, figs. 6, 7). The en- 
docarp composed of interdigitating rounded cells, and 
exocarp of thin-walled rectangular cells are distinctive 
features of this species (Pl. 67, fig. 8). The unilocular, 
single-seeded morphology and interconnecting cells of 
the endocarp are reminiscent of Icacinaceae; however, 
the endocarp shows no obvious plane of dehiscence as 
occurs in many Icacinaceae and a funicular canal was 
not detected. The affinities of this species, whether 
within the Icacinaceae, or with another family, remain 
to be determined. 


Carpolithus sp. 6 
Plate 67, figures 9-12 


Description.—Fruit subovoid, unilocular, single- 
seeded, base and apex rounded, length 5.1 mm, width 
4.2 mm, thickness 3.9 mm; endocarp ellipsoidal, bi- 
symmetrical, laterally compressed with a keel in the 
plane of symmetry, endocarp wall 0.2 mm thick, with 
smooth locule and verrucate outer surface, composed 
of thick-walled sclereids 20-38 um in diameter; me- 
socarp 0.6—0.9 mm thick, composed of large, polygo- 
nal, isodiametric to anticlinally elongate parenchyma 
cells 80-160 um in diameter, these cells smaller in the 
outer few peripheral layers and with about 30 longi- 
tudinally oriented vascular bundles close to the en- 
docarp; large empty cavities developed in parts of the 
mesocarp; placentation ventral near the base; seed fill- 
ing the locule, seed coat composed of two to three layers 
of rectangular, periclinally elongate cells; raphe with 
several vascular bundles forming a band 400 um thick 
and 100 um wide along one of the lateral margins 
adjacent to the keel of the endocarp. 

Specimen. — USNM 424738. 

Discussion.— This species is known from a single 
specimen that is not very distinctive in its surface mor- 
phology (Pl. 67, figs. 9, 10), but in section shows good 
anatomical structure (Pl. 67, figs. 11, 12). The meso- 
carp appears to have been rather soft, and one would 
expect the endocarp alone to be more commonly pre- 
served. However, I have not seen corresponding iso- 
lated endocarps with the distinctive verrucate surface 
ornamentation observed in this species. 


Carpolithus sp. 7 
Plate 67, figures 13-16 


Description.— Locule cast subglobose-oblate, round- 
ed basally, truncate apically, height 7.5, 11, 12.1 mm, 
equatorial diameter 8.5, 13.8, 14.1 mm; surface smooth 
to irregularly wrinkled; truncation smooth, more or 
less circular, perpendicular to the vertical axis, its mar- 


gins somewhat recessed, slightly domed (wide-obtusely 
conical) forming an obtuse point in the center. Mold 
in matrix globose, with very little space between cast 
and mold, at most 0.2 mm, with a small domed pro- 
jection at one end. 

Specimens.—UF 9862, USNM 355191, 355637. 

Discussion.— This species is distinctive in its globose 
shape and circular, shallowly domed truncation. One 
of the specimens (UF 9862) is a cast along with a mold 
of the surrounding endocarp. The space between the 
locule cast and the endocarp mold, representing the 
endocarp wall thickness, is only about 0.2 mm. Such 
a thin wall may explain the surface wrinkling, which 
could be due to compaction in the sediment. 


Carpolithus sp. 8 
Plate 68, figures 1-4 


Description. — Achene more or less ovoid, truncate 
at one end, rounded-acute at the other; more or less 
circular in cross section, length 4.0 mm, equatorial 
diameter 3.4 mm, exterior smooth, base with a ге- 
cessed circular scar, wall relatively thick (0.8 mm), 
composed of isodiametric cells 23—63 um in diameter; 
locule lining composed of a single layer of small cu- 
boidal cells ca. 7 um in diameter. 

Specimen.— USNM 422526. 

Discussion.—' This species is known from a single 
specimen that is distinctive in its thick wall and large 
basal scar. Preliminary study indicates close similarity 
in morphology and anatomy to extant Sparganium. 
Sparganium achenes are truncate at the apex, and, like 
those of the fossils, have an endocarp of a single rel- 
atively thick layer of isodiametric cells. 


Carpolithus sp. 9 
Plate 68, figures 5-9 


Description.— Fruit or bud subglobose, trigonal in 
cross section, pointed at one end, truncate at the other, 
with three longitudinal keels intersecting at the pointed 
end; length 2.0-2.5 mm, width 1.6-2.4 mm; truncate 
end with a central circular scar (style base or peduncle 
scar?) surrounded by four alternately arranged whorls 
of three slits interpreted as tepal scars; wall (endocarp?) 
200 um thick. 

Specimens.—UF 6347, 6457, 9550-9556. | 

Discussion. — These specimens are distinctive in their 


trigonal symmetry and truncated end with whorls of 


slits that appear to represent tepals. Whether these 
objects represent immature buds or actual fruits has 
not been determined. Transverse sections failed to ге“ 
veal stamens, as might be expected if the specimens 
represent flower buds. The sections also did not reveal 
seeds. 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 119 


Carpolithus sp. 10 
Plate 68, figures 10-14 


Description.— Fruits/seeds trigonal in face view, bi- 
Symmetrical, with a dorsal keel in plane of symmetry, 
height 1.4-1.7, аур. 1.6 mm, (SD=0.13, n=4), width 
across plane of symmetry 0.9-1.1, avg. 1.0 mm 
(SD=0.08, n=4), depth in plane of symmetry 1.6-1.7, 
avg. 1.7 mm (SD=0.06, n=3); rounded dorsally, nar- 


rowly triangular in ventral view; with pore (hilum or ` 


funicular opening) circular on the ventral surface. 

Specimens.—USNM 435102-435105. 

Discussion.—This species is represented only by a 
few specimens. In shape and size they compare favor- 
ably with Brousonettia in the Moraceae. The species 
of Brousonettia that I have seen have a verrucate seed 
Coat, whereas this fossil species has a smooth outer 
Surface. Moraceous wood anatomically similar to 
Brousonettia is present among the fossil woods from 
the Nut Beds (Manchester, unpublished). 


Carpolithus sp. 11 
Plate 68, figures 15-19 


Etymology.—The epithet refers to the Clarno For- 
Mation. 

Description.—Inner mold of seed coat oblong in face 
View, bilaterally symmetrical; base rounded, apex ap- 
Parently rounded, dorsiventrally compressed, some- 
what curved ventrally from base to apex as seen from 
the side, length ca. 11.3 mm, width ca. 4.9 mm, thick- 
Dess ca. 2.0 mm, with a median keel on the dorsal 
Surface and broad median depression on the ventral 
Side; surface of mold with about 22 closely spaced, 
Tounded horizontal ribs arising oppositely from the 
Median keel. These ribs are straight in course, passing 
from the dorsal keel over the dorsal face and around 
to the ventral side where the meet in an offset pattern, 
the ribs perpendicular to the long axis of the seed except 
at the base and apex where they are radially arranged. 

Specimen.—USNM 424625. 

Discussion.—This species is represented by a single 
very distinctive specimen showing a series of rounded 
transverse ridges separated by narrow cleavages (PI. 
68, figs. 15—19). The specimen is interpreted as the 
Internal mold of a seed, suggesting that the seed coat 
Itself had a series of transverse partial septa and that 
the endosperm was therefore ruminate. Unlike the seeds 
of Annonaceae with ruminate endosperm (PI. 6, figs. 
9-18), the species has a prominent dorsal keel, and the 
ribs are more parallel and even in their development. 
Ruminate endosperm also occurs among the Menis- 
Permaceae (Thanikaimoni, 1984), and in shape and 
Symmetry the fossil resembles endocarps of the Ti- 
Nosporeae tribe; however, I have not found a closely 


similar modern genus, and the apex of the fossil is 
broken, making it impossible to determine if placen- 
tation was subapical as in Tinosporeae. 


Carpolithus sp. 12 
Plate 69, figure 1-4 


Description.—Fruit a penta-syncarpellate capsule, 
prolate, rounded in lateral view, with five locules ra- 
dially arranged about a central axis; length 11.5 mm, 
width 8.3-10.1 mm; surface smooth, each carpel with 
a median longitudinal dorsal keel, possibly a line of 
dehiscence; adjacent carpels separated by a groove on 
the external surface and by a thin septum (ca. 0.4 mm) 
formed by the paired carpel walls; number and type 
of seeds unknown. 

Specimen.— USNM 326719 (Bones, 1979, pl. 5, fig. 
10). 

Discussion. — This species is represented by a single 
specimen with five conspicuously bulging locules (Pl. 
69, figs. 1-3). The fruit was sectioned transversely re- 
vealing thin septa, a thick central axis, and empty loc- 
ules (Pl. 69, fig. 4) suggesting that the seeds had dis- 
persed prior to deposition of the fruit. 


Carpolithus sp. 13 
Plate 69, figures 5-8 


Description. —Fruit subglobose, unilocular, single- 
seeded?, rounded apically and basally; length and width 
15.0 mm; surface smooth; wall 2.7 mm thick, com- 
posed of: locule lining 6-7-seriate, 600 um thick, com- 
posed of anticlinally elongate cells 70-120 um long, 
20-45 um wide; outside the locule lining is a uniseriate 
layer 20 um thick of narrow anticlinally elongate cells, 
poorly preserved, frequently expanded and containing 
rhomboidal crystals; mesocarp not preserved, repre- 
sented by a space 1.6 mm thick penetrated by foreign 
roots; exocarp composed of a layer of isodiametric 
sclereids 50-75 um in diameter, outlined by a unise- 
riate layer 15 um thick composed of rectangular cells 
15-30 um in diameter. 

Specimen. —UF 9744. 

Discussion.— This species is represented by a single 
specimen. Although the exterior of the specimen is 
fractured and partially covered with sediment (Pl. 69, 
fig. 5), the internal structure and anatomy is very well 
preserved as seen in transverse sections (Pl. 69, figs. 
6-8). The inner layer of short, anticlinally elongate 
thin-walled cells (Pl. 69, fig. 7) and the outer layer of 
compact isodiametric sclereids (Pl. 69, figs. 8) are con- 
spicuous features of this fruit. 


Carpolithus sp. 14 
Plate 69, figures 9-12 


Description.—Fruit a multiseeded berry, pyriform, 


120 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


circular in cross section, pointed apically, rounded with 
a slight depression and 1 mm circular scar at the base; 
length 12.2 mm, equatorial diameter 11.3 mm; fruit 
surface smooth, evidently thin-walled; placentation 
type uncertain, seeds ovoid, 1.8-2.8 wide, ca. 5.0 mm 
long, borne in three pairs near the center of the fruit 
and surrounded by pulpy tissue; seed coat composed 
оГа layer of uniseriate, anticlinally oriented columnar 
cells 50-70 um high and 20-25 um wide. 

Specimen.—UM 66137. 

Discussion. — This species is known only from a sin- 
gle specimen. Transverse sectioning showed that much 
of the inner contents had degraded prior to silicifica- 
tion; however, a central group of six paired seeds are 
recognizable, two of which are well preserved (Pl. 69, 
figs. 11, 12). Toward the periphery of the fruit are 
remains of what appear to be degraded parenchyma 
tissue suggesting that it was a fleshy fruit. 


Carpolithus sp. 15 
Plate 69, figures 13-16 


Description.— Cast bisymmetrical, H-shaped in face 
view, consisting of two crescent shaped limbs, each 
more or less parallel, concave ventrally and convex 
dorsally, rounded at one end, narrowed and pointed 
at the other; these limbs mutually joined connected 
across the plane of symmetry in the central % of their 
length; height 5.7 mm, width across plane of symmetry 
4.0 mm, thickness 3.5 mm; surface nearly smooth, but 
finely striate by surface cells oriented horizontally with 
respect to the long axis ofeach limb; dorsiventral thick- 
ness of each limb 2.2 mm, each limb with a dorsal keel 
that is best developed at the narrow end. 

Specimen.—UF 9548. 

Discussion.— Only a single specimen of this type is 
known. It is distinctive in the H-shape in face view 
(Pl. 69, figs. 13, 14) and strongly curved longitudinal 
axis as viewed laterally (Pl. 69, fig. 15). The structure 
apparently represents the inner mold ofa seed (embryo 
cast) or fruit (locule cast). If it is an embryo cast then 
the two limbs might be interpreted as a pair of thick 
cotyledons. If it is a locule cast, then the fruit appears 
to have been almost bilocular, but with a foramen 
between the two lateral halves. 


Carpolithus sp. 16 
Plate 70, figures 1-5 


Description.—Seed laterally flattened, ellipsoidal, bi- 
laterally symmetrical with a ventral keel in plane of 
symmetry, rounded dorsally, length 3.2 mm, width 2.0 
mm, thickness 1—1.2 mm; surface with prominent nar- 
row rugulations perpendicular to long axis of seed. Em- 
bryo cavity smooth-surfaced, laterally compressed like 
the seed, roughly J-shaped with notch in the placental 
area. 


Specimens.— UF 9609, 9794-9798. 

Discussion.—Seeds of this species are preserved to- 
gether in clusters within the siltstone. Typically the 
seed coat itself has deteriorated, leaving a rugulate out- 
er mold in the siltstone with a silica cast of the embryo 
cavity inside. The surficial relief on the seeds can be 
seen through stereo images of the matrix mold (PI. 70, 
figs. 1, 2), and on silicone casts prepared from the same 
specimens (Pl. 70, fig. 5). Although internal compo- 
sition of the seed is unknown, internal molds of the 
seed coat (Pl. 70, figs. 3, 4) are curved, suggesting а 
curved embryo, with an attachment scar at the axis of 
curvature. 


Carpolithus sp. 17 
Plate 70, figures 6-8 


Description.— Fruit globose-prolate, unilocular, sin- 
gle-seeded, base and apex rounded, circular in trans- 
verse section, length 6.2 mm, equatorial diameter 515 
mm; pericarp 1.25 mm thick, consisting of a thick 
parenchymatous layer of isodiametric cells 30-70 шт 
in diameter surrounded by a uniseriate layer of anti- 
clinally elongate schlerenchyma cells 210 um high, 30- 
38 um wide; seed coat 38 um thick, four- to six-seriate, 
composed ofthin walled polygonal cells 8-15 um high, 
12-30 um, seed filled with parenchyma 50-180 um in 
diameter. 

Specimen.— UF 9768. 

Discussion.— This globose unilocular specimen shows 
some similarity to the fruits of Lauraceae. However, 
although it has an outer layer of columnar cells, its 
lacks such a layer close to the locule and it does not 
show the large, subhemispherical cotyledons typical of 
Lauraceous seeds. 


Carpolithus sp. 18 
Plate 70, figures 9-12 


Description.—Fruit broadly elliptical in face view, 
lenticular to compressed-hexagonal in cross section, 
unilocular, quadrilaterally symmetrical with the two 
planes of symmetry intersecting at right angles in the 
longitudinal axis), with a median keel in the dorsiven- 
tral plane of symmetry; ventral and dorsal faces round- 
ed or with a pair of longitudinal angles, base rounded- 
truncate with a persistent stalk 0.7 mm long and 0.4 
mm thick, apex rounded; fruit length 6.8 mm, width 
6.2 mm, thickness 4.7 mm; wall 123-150 um thick, 
seed similar in shape to the fruit, raphe running lon- 
gitudinally down one of the narrow sides. 

Specimens.—UF 6497, 9464. 

Discussion.— This species is represented only by two 
specimens, one of which is fragmentary. One of them 
was sectioned transversely, revealing a thin wall and 
large locule devoid of contents (Pl. 70, fig. 12). In the 


| 
| 
| 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 121 


Cutting process, Ше wall flaked away on Ше apical half, 
revealing an internal cast of the seed showing the course 
of the raphe on one side along the keel. The description 
above uses terminology that assumes the specimen is 
а, fruit containing a single seed. Alternatively, the spec- 
Imen might be interpreted as a seed with the outer wall 
representing the seed coat and the projection а prom- 
Inent protruding hilum. The other specimen shows 


Some poorly preserved dark organic material adhering _ 


to the fruit cast, indicating that the specimens are miss- 
Ing the outer layer(s). 


Carpolithus sp. 19 
Plate 70, figures 13, 14 


Description.—Cast globose, rounded apically and 
basally, length 15.2 mm, equatorial diameter 15.6 mm; 
apical l^ with three equidistant faint longitudinal ridges 
Joining at the apex to form a triradiate mark; surface 
Smooth in the lower hemisphere, striate in the upper 
hemisphere with longitudinally aligned low rounded 
ribs that are intertwining and plicate at their extrem- 
Ities, 

Specimen. — USNM 422381. 

Discussion.— This specimen is a cast in translucent 
Chalcedony without cellular preservation. It appears to 
be the internal mold ofa fibrous endocarp or seed coat. 
Because of the triradiate mark at the apex (Pl. 70, figs. 
13, 14), it brings to mind Palmae. There is no evidence, 
however, of germination openings. 


Carpolithus sp. 20 
Plate 70, figures 15, 16 


Description.— Fruit globose, unilocular, single-seed- 
ed, rounded apically and basally, length 7.5 mm, width 
6.5 mm; wall 0.9 mm thick, composed of isodiametric 
Sclereids 38—63 um in diameter. 

Specimens. —UF 9759, 9760. 

Discussion.— This species is represented by two 


specimens. The layer of isodiametric sclereids com- 
posing the fruit wall distinguishes it from other uni- 
locular fruits of the Nut Beds. Although similar in size 
and shape to Prunus weinsteinii, this species does not 
show a keel or plane of dehiscence. 


Carpolithus sp. 21 
Plate 70, figure 18 


Description.—Seed cordate, quadrilaterally sym- 
metrical, length 6.5 mm, width 8 mm, thickness 3.8 
mm (est.); internal mold of seed coat more or less 
triangular in face view, dorsiventrally compressed, fu- 
siform in transverse section, smooth to faintly rugulate, 
5.8 mm long, 5.7 mm wide, 1.9 mm thick, rounded 
basally, with a protruding scar of attachment, pointed 
apically; seed coat thickness 1.2 mm. 

Specimen.— USNM 422525. 

Discussion.— This species is known only from a sin- 
gle specimen. Although it bears some resemblance to 
seeds of Magnoliaceae, it is perfectly symmetrical. The 
inner cast is smooth, however the surrounding mold 
in the sediment is rough-textured with protrusions (Pl. 
70, fig. 18). 


Five-part flower 
Plate 70, figure 17 


Description.— Flower 5-parted, actinomorphic, ca. 
18 mm wide, with central circular structure (ovary?) 
4.5 mm in diameter, five free ovate sepals 7.0 mm 
long, 3-5 mm wide, five free petals alternate with and 
thinner than the sepals, with pronounced parallel veins. 
Pedicel 1.2 mm in diameter, at least 3 mm long. 

Specimens. — OMSI Pb1507, 1642. 

Discussion. — Two flowers of this type were recov- 
ered from the leaf stratum at the base of Face 3. One 
is transversely compressed (Pl. 70, fig. 17), the other 
is laterally compressed. 


REFERENCES CITED 


Airy Shaw, H. K., ed. 

1973. A dictionary of the flowering plants and ferns. 8th ed. Cam- 

bridge University Press, Cambridge, England, 1245 pp. 
Axelrod, D. L, Arroyo, M. T., and Raven, P. Н. 

1991. Historical development of temperate vegetation in the 
Americas. Revisita Chilena de Historia Natural, vol. 64, 
pp. 413-446. 

Bailey, I. W. and Howard, R. A. 

1941. The comparative morphology of the Icacinaceae I. Anat- 
omy of the node and internode. Journal of the Arnold 
Arboretum, vol. 22, pp. 125-132. 

Basinger, J. F. 

1981. The vegetative body of Metasequoia milleri from the Mid- 
dle Eocene of southern British Columbia. Canadian Jour- 
nal of Botany, vol. 59, pp. 2379-2410. 


Basinger, J. F. and Rothwell, G. W. 
1977. Anatomically preserved plants from the Middle Eocene (Al- 
lenby Formation) of British Columbia. Canadian Journal 
of Botany, vol. 55, pp. 1984-1990. 
Becker, H. F. 
1969. Fossil plants of the Tertiary Beaverhead Basins in south- 
western Montana. Palaeontographica, Abt. B, vol. 127, 
pp. 1-142, pl. 1-44. 
Berry, E. W. 
1930. Revision of the Lower Eocene Wilcox flora of the south- 
eastern United States. U.S. Geological Survey Professional 
Paper 156, pp. 1-196. 
Bestland, E. A., Retallack, G. J., Swisher, C. C., and Fremd, T. J. 
1993. Timing of cut-and-fill sequences in the John Day For- 
mation (Eocene-Oligocene), Painted Hills Area, Central 


122 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Oregon. Geological Society of America, Abstracts with Pro- 
grams, vol..25,.n0.-5, p. 9. 
Beusekom, C. F. van 
1971. Revision of Meliosma (Sabiaceae), section Lorenzanea ex- 
cepted, living and fossil, geography and phylogeny. Blumea, 
vol. 19, pp. 355-529. 
Blanc-Louvel, C. ; 
1985. Paléoflore du Gisement Eocene de Prémontré dans L Aisne: 
Étude de deux fruits. 110* Congrés national des Sociétés 
savantes, Montpellier, 1985, sciences, fasc. 5, pp. 149- 
161, pl. 1-3. 
Bogle, A. L. and Philbrick, С. Т. 
1980. A generic atlas of hamamelidaceous pollens. Contributions 
of the Gray Herbarium, vol. 210, pp. 29-103. 
Bones, T. J. 
1979. Atlas of fossil fruits and seeds from North Central Oregon. 
Oregon Museum of Science and Industry Occasional Pa- 
pers in Natural Science, vol. 1, pp. 3-23. 
Bremekamp, C. E. B. 
1966. Remarks on the position, the delimitation and the subdi- 
vision of the Rubiaceae. Acta Botanica Neerlandica, vol. 
15, pp. 1-33. 
Bremer, B. and Struwe, L. 
1992. Phylogeny of the Rubiaceae and the Loganiaceae: Con- 
gruence or conflict between morphological and molecular 
data. American Journal of Botany, vol. 79, pp. 1171- 
1184. 
Brown, J. Т. 
1975. Equisetum clarnoi, a new species based on petrifactions 
from the Eocene of Oregon. American Journal of Botany, 
vol. 62, pp. 410-415. 
Brown, R. W. 
1929. Additions to the flora of the Green River Formation. U.S. 
Geological Survey Professional Paper 154-J, pp. 279-293, 
7 ple 
1940. New species and changes of name in some American fossil 
floras. Journal of the Washington Academy of Sciences, 
vol. 30, pp. 344-353. 
Camus, A. 
1952-1954. Les chénes. in Encylopédie Economique de Sylvi- 
culture, vol. 8, pp. 511-1196. 
Cevallos-Ferriz, S. R. S. and Stockey, R. A. 
1988a. Permineralized fruits and seeds from the Princeton chert 
(Middle Eocene) of British Columbia: Lythraceae. Cana- 
dian Journal of Botany, vol. 66, pp. 303-312. 
1988b. Permineralized fruits and seeds from the Princeton chert 
(Middle Eocene of British Columbia: Araceae. American 
Journal of Botany, vol. 75, pp. 1099-1113. 
1989. Permineralized fruits and seeds from the Princeton chert 
(Middle Eocene) of British Columbia: Nymphaeaceae. Bo- 
tanical Gazette, vol. 150, pp. 207-217. 
1990. Permineralized fruits and seeds from the Princeton chert 
(Middle Eocene) of British Columbia: Vitaceae. Canadian 
Journal of Botany, vol. 68, pp. 288-295. 
1991. Fruits and seeds from the Princeton chert (Middle Eocene) 
of British Columbia: Rosaceae (Prunoideae). Botanical Ga- 
zette, vol. 152, pp. 369-379. 
Chandler, M. E. J. 
1926. The Upper Eocene flora of Hordle, Hants. The Palaeon- 
tographical Society, London, 52 pp., 8 pl. 
1957. The Oligocene flora of the Bovey Tracey Lake Basin, De- 
vonshire. Bulletin of the British Museum (Natural History) 
Geology, vol. 3, pp. 71-123, pl. 11-17. 
1960. Plant remains of Hengistbury and Barton Beds. Bulletin 


of the British Museum (Natural History) Geology, vol. 4, 
pp. 193-238, pl. 29-35. 

1961a. Flora of the Lower Headon Beds of Hampshire and the 
Isle of Wight. Bulletin of the British Museum (Natural 
History) Geology, vol. 5, pp. 91-158, pl. 24-30. 

1961b. The Lower Tertiary floras of southern England. 1. Paleo- 
cene Floras. London Clay Flora (Supplement). Text and 
Atlas. British Museum (Natural History), London, xi + 
354 pp., 34 pl. 

1962. The Lower Tertiary floras of southern England. 2. Flora 
ofthe Pipe-Clay Series of Dorset (Lower Bagshot). British 
Museum (Natural History), London, xi + 176 pp., 29 pl. 

1963a. Revision of the Oligocene floras of the Isle of Wight. Bul- 
letin of the British Museum (Natural History) Geology, 
vol. 6, pp. 321-384, pl. 27-35. 

1963b. The Lower Tertiary floras of southern England. 3. Flora 
of the Bournemouth Beds; the Boscombe, and the H ighcliff 
Sands. British Museum (Natural History), London, xi + 
169 pp., 25 pl. 

1964. Thelower Tertiary floras of southern England. 4. Summary 
and survey of findings in the light of recent botanical ob- 
servations. British Museum (Natural History), London, xii 
+ 151 pp., 4 pl. 

1978. Supplement to the Lower Tertiary Floras of Southern En- 
gland. Tertiary Research Special Paper, vol. 4, pp. 1-44. 

Chaney, R. W. 

1927. Geology and Palaeontology of the Crooked River Basin, 
with special reference to the Bridge Creek flora. Carnegie 
Institute of Washington Publication 346, pp. 45-138, pl. 
2-20. 

Cheesman, Е. Е. 

1947. Classification ofthe bananas. I. The genus Ensete Horan. 

Kew Bulletin, vol. 2, pp. 97-117. 
Chernik, V.V. 

1980. Peculiarities of structure and development of the pericarp 
of the representatives of the family Ulmaceae and Celti- 
daceae. Botanicheskii Zhurnal, vol. 65, pp. 521-531 (In 
Russian). 

Collinson, M. E. 

1982. А preliminary report on the Senckenberg-Museum collec- 
tion of fruits and seeds from Messel bei Darmstadt. Courier 
Forschungsinstitut Senckenberg, vol. 56, pp. 49-57. 

1983. Fossil plants of the London Clay. The Palaeontological 
Association, London, England, 121 pp. 

1988. The special significance of the Middle Eocene fruit and seed 
flora from Messel, Western Germany. Courier Forschung- 
sinstitut Senckenberg, vol. 107, pp. 187-197. 

Collinson, M. E. and Hooker, J. J. 

1987. Vegetational and mammalian faunal changes in the Early 
Tertiary of southern England, in Friis, E. M., Chaloner, 
W. G., and Crane, Р. R. [eds.], The origins of angiosperms 
and their biological consequences, pp. 259-304. Cam- 
bridge University Press, Cambridge, England. 

1992. Fossil evidence of interactions between plants and plant- 
eating mammals. Philosophical Transactions of the Royal 
Society, London, В, vol. 333, pp. 197—208. 

Corner, E. J. H. 

1976. The seeds of dicotyledons. Cambridge University Press, 

Cambridge, England, 2 vol., 311 + 552 pp. 
Cowan, D. 5. 

1982. Geological evidence for a post-40 m.y. B.P. large-scale dis- 
placement of part of southeastern Alaska. Geology, vol. 
10, pp. 309-313. 


| 
| 
| 
| 
| 
| 
| 
| 
| 


| 
| 
| 
| 
| 
| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 123 


Crane, P.R. 

1981. Betulaceous leaves and fruits from the British Upper Pa- 
laeocene. Botanical Journal of the Linnean Society, vol. 
83, pp. 103-136. 

1987. Vegetational consequences of the angiosperm diversifica- 
tion, in Friis, E. M., Chaloner, W. G., and Crane, Р. К. 
[eds.], The origins of angiosperms and their biological con- 
sequences. pp. 107-144, Cambridge University Press, 
Cambridge, England. 

1989. Early fossil history and evolution of the Betulaceae, in Crane, 


Р. К. and Blackmore, 5. [eds.], Evolution, Systematics, and · 


fossil history of the Hamamelidae, Volume 2: ‘Higher’ 

Hamamelidae. Systematics Association Special Volume 

no. 40B, pp. 87-116, Clarendon Press, Oxford, England. 
Crane, P. R., Manchester, S. R., and Dilcher, D. L. 

1990. A preliminary survey of fossil leaves and well-preserved 
reprod structures from the Sentinel Butte Formation (Pa- 
leocene) near Almont, North Dakota. Fieldiana Geology, 
New Series, no. 20, pp. 1-63. 

Crepet, W. L. 

1989. History and implications of the early North American rec- 
ord of Fagaceae, in Crane, P. R. and Blackmore, S. [eds.]. 
Evolution, Systematics, and fossil history of the Hama- 
melidae, Volume 2: “Higher” Hamamelidae. Systematics 
Association Special Volume no. 40B, pp. 45-66, Claren- 
don Press, Oxford, England. 

Crepet, W. L. and Nixon, К. С. 

1989a. Earliest megafossil evidence of Fagaceae: phylogenetic and 
biogeographic implications. American Journal of Botany, 
vol. 76, pp. 842-855. 

1989b. Extinct transitional Fagaceae from the Oligocene and their 
phylogenetic implications. American Journal of Botany, 
vol. 76, pp. 1493-1505. 

Cronquist, A. 

1981. An integrated system of classification of flowering plants. 

Columbia University Press, New York, 1262 pp. 
Czeczott, Н. and Skirgiello, A. 

1959. The fossil flora of Turow near Bogatynia I and II (1), 
Dicotyledones. Prace Muzeum Ziemi, vol. 3, pp. 94-112, 
121-128. 

Daghlian, C. P. 

1978. Coryphoid palms from the lower and middle Eocene of 
southeastern North America. Palaeontographica, Abt. B, 
vol. 166, pp. 44-82, 15 pl. 

Daghlian, С. Р. and Crepet, W. L. 

1983. Oak catkins, leaves and fruits from the Oligocene Cata- 
houla Formation and their evolutionary significance. 
American Journal of Botany, vol. 70, pp. 639-649. 

Daly, D. C. 

1989. Studies in neotropical Burseraceae. II. Generic limits in 
New World Protieae and Canarieae. Brittonia, vol. 4, pp. 
17-27. 

Diels, L. 
1910. Menispermaceae. Das Pflanzenreich, vol. 46 (IV-94), 345 


pp. 
Dilcher, D. L. 
1971. A revision of the Eocene flora of southeastern North Amer- 
ica. The Paleobotanist, vol. 20, pp. 7-18. 
Dilcher, D. L., and McQuade, J. F. 
1967. A morphological study of Nyssa endocarps from Eocene 
deposits in western Tennessee. Bulletin of the Torrey Bo- 
tanical Club, vol. 94, pp. 35-40. 
Dorofeev, PE 
1963. The Tertiary floras of western Siberia. Akademia Nauk 


S.S.S.R., Moscow, 345 pp., 50 pl. (in Russian with English 
Summary). 

1982. Species determined from fruits. Celtis and Aphananthe, in 
Takhtajan, A. and Zhilin, S. [eds.], Fossil flowering plants 
of the USSR, vol. 2, Ulmaceae— Betulaceae. pp. 22-24, 
Nauka, Leningrad. (in Russian). 

Doyle, J. A., Schorn, H. E., Tiffney, B. H., and Upchurch, G. R., 
Jr. 

1988. The LaPorte flora-earliest Oligocene of north-central Cal- 
ifornia. Field Guide Book for the 1988 meeting of the 
Paleobotanical Section of the Botanical Society of Amer- 
ica, Davis, California, August 14, 1988. 

Drinnan, А. N., Crane, P. R., Friis, E. M., and Pedersen, К. К. 

1990. Lauraceous flowers from the Potomac Group (mid-Cre- 
taceous) of eastern North America. Botanical Gazette, vol. 
151, pp. 370-384. 

Dupler, A. W. 

1920. Ovuliferous structures of Taxus canadensis. Botanical Ga- 

zette, vol. 69, pp. 492-519. 
Endress, P. K. 

1989a. A suprageneric taxonomic classification of the Hamame- 
lidaceae. Taxon, vol. 38, pp. 371-376. 

1989b. Aspects of evolutionary differentiation of the Hamameli- 
daceae and the Lower Hamamelididae. Plant Systematics 
and Evolution, vol. 162, pp. 193-211. 

1989c. Phylogenetic relationships in the Hamamelidoideae, in 
Crane, P. R. and Blackmore, S. [eds.]. Evolution, System- 
atics, and fossil history of the Hamamelidae, Volume 1: 
*Lower Hamamelidae. Systematics Association Special 
Volume no. 40B, pp. 227-248, Clarendon Press, Oxford, 
England. 

Endress, P. K. and Friis, E. M. 

1990. Archamamelis, hamamelidalean flowers from the Upper 
Cretaceous of Sweden. Plant Systematics and Evolution, 
vol. 175, pp. 101-114. 

Engler, A. 

1897. Icacinaceae. Ш, Abt. 5. Die natürlichen Pflanzenfamilien, 
] aufl. Leipzig. 

1964. Syllabus der pflanzenfamilien. 12 ed., Vol. 2. Angiosper- 
men übersicht über die Florengebiete der Erde. Gebrüder 
Borntraeger, Berlin. 

Erwin, D. M. E 

1987. Silicified palm remains from the Middle Eocene (Allenby 
Formation) of British Columbia [abs.]. American Journal 
of Botany, vol. 74, p. 681. 

Evernden, J. F., Savage, D. E., Curtis, G. H., and James, G. T. 

1964. Potassium-argon dates and the Cenozoic mammalian 
chronology of North America. American Journal of Sci- 
ence, vol. 262, pp. 145-198. 

Eyde, R. H. ; 

1963. Morphological and paleobotanical studies of the Nyssa- 
ceae, I. a survey of the modern species and their fruits. 
Journal of the Arnold Arboretum, vol. 44, pp. 1-54, 5 pl. 

1968. Flowers, fruits, and phylogeny of the Alangiaceae. Journal 
of the Arnold Arboretum, vol. 49, pp. 167-192. 

1988. Comprehending Cornus: Puzzels, and progress in the sys- 
tematics of the dogwoods. Botanical Review, vol. 54, pp. 
233-351. 

Eyde, R. H., and Barghoorn, E. S. 

1963. Morphological and paleobotanical studies of the Nyssa- 
ceae, II: the fossil record. Journal of the Arnold Arbore- 
tum, vol. 44, pp. 328-370. 


124 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Eyde, К. H., Bartlett, A., and Barghoorn, К. 5. 

1969. Fossil record of Alangium. Bulletin of the Torrey Botanical 

Club, vol. 96, pp. 288-314. 
Eyde, R. H. and Q., Xiang. 

1990. Fossil mastixioid (Cornaceae) alive in eastern Asia. Amer- 
ican Journal of Botany, vol. 77, pp. 689-692. 
Fiebelkorn, R. B., Walker, С. W., MacLeod, N. S., McKee, E. H., 

and Smith, J. G. 

1983. Index to K-Ar age determinations for the State of Oregon. 
Isocron/West, vol. 37, pp. 1-60. 

Forman, L. L. 

1966. Generic delimitation in the Castaneoideae. Kew Bulletin, 
vol. 18, pp. 421-426. 

1974. The endocarps of Cocculus (Menispermaceae). Kew Bul- 
letin, vol. 29, pp. 477-481. 

1985. A revision of the tribe Fibraureae (Menispermaceae) in 
Asia. The Mensipermaceae of Malesia and adjacent areas 
XIII. Kew Bulletin, vol. 40, pp. 539-551. 

1986. Menispermaceae. Flora Malesiana, ser. I, vol. 102, pp. 
157-253. 

Foster, S. A. 

1986. On the adaptive value of large seeds for tropical moist forest 
trees: a review and synthesis. Botanical Review, vol. 52, 
pp. 260-299. 

Foster, S. A. and Janson, C. H. 

1985. The relationship between seed size and establishment con- 
ditions in tropical woody plants. Ecology, vol. 66, pp. 773— 
780. 

Friis, E. M. 

1979. The Damgaard flora: a new Middle Miocene flora from 
Denmark. Bulletin of the Geological Society of Denmark, 
vol. 27, pp. 117-142. 

1985. Angiosperm fruits and seeds from the Middle Miocene of 
Jutland (Denmark). Biologiske Skrifter Det Kongelige 
Danske Videnskaberne Selskab, vol. 24, pp. 1-165. 

Friis, E. M., Crane, P. R., and Pedersen, K. R. 

1988. Reproductive structures of Cretaceous Platanaceae. The 
Royal Danish Academy of Sciences and Letters Biologiske 
Skrifter, vol. 31, pp. 1-55. 

Geissert, F. and Gregor, H.-J. 

1981. Einige interessante und neue sommergrüne Pflanzenele- 
mente (Fruktifikationen) aus dem Elsässer Pliozan (Sabia 
Colebr., Wikstroemia Endl., Alangium Lam., Nyssa L., 
Halesia Ellis, Rehderodendron Hu). Mitteilungen des bad- 
ischen Landesvereins Naturkunde und Naturschutz, vol. 
12, pp. 233-239. 

Gentry, A. Н. 

1983. Dispersal ecology and diversity in Neotropical forest com- 
munities. Sonderband Naturwissenshaftlichen Vereins in 
Hamburg, vol. 7, pp. 303-314. 

1992. The distribution and evolution of climbing plants, in F. E. 
Putz, and Mooney, H. A. [eds.], The biology of vines, pp. 
3-43, Cambridge University Press, New York. 

Gregor, H.-J. 

1978. Die miozänen Frucht-und Samen-Floren der Oberpfälzer 
Braunkohle. I. Funde aus den sandigen Zwischenmitteln. 
Palaeontographica, Abt. B, vol. 167, pp. 1-103. 

1978. Schizandra geissertii nova spec.—ein exotisches Element 
im Elsässer Pliozän (Sessenheim, Brunssumien). Mittei- 
lungen des badischen Landesvereins Naturkunde und Na- 
turschutz, vol. 12, pp. 241-247. 

Gregory, I. 

1969. Fossilized palm wood in Oregon. Ore Bin, vol. 31, pp. 93- 

110. 


Greuter, W., editorial chairman. 

1988. International Code of Botanical Nomenclature. Adopted 
by the 14th International Botanical Congress, Berlin, Koeltz 
Scientific Books, Konigstein, 328 pp. 

Grote, P. J. 

1989. Selected fruits and seeds from the Middle Eocene Claiborne 
Formation of southeastern North America. PhD disserta- 
tion, Indiana University, Bloomington, Indiana 233 pp. 

Grote, P. J. and Dilcher, D. L. 

1989. Investigations of angiosperms from the Eocene of North 
America: a new genus of Theaceae based on fruit and seed 
remains. Botanical Gazette, vol. 150, pp. 190-206. 

Grudzinskaya, I. A. 

1967. Ulmaceae and reasons for distinguishing Celtidaceae as а 
separate family Celtidaceae Link. Botanicheskii Zhurnal, 
vol. 52, pp. 1723-1748. 

Gunn, C. R. 

1984. Fruits and seeds of genera in the subfamily Mimosoideae 
(Fabaceae). U.S. Department of Agriculture, Agricultural 
Research Service, Technical Bulletin 1861, 194 pp. 

1991. Fruits and seeds of genera in the subfamily Caesalpinioi- 
deae (Fabaceae). U.S. Department of Agriculture, Agri- 
cultural Research Service, Technical Bulletin 1755, 408 
рр. 

Hammel, В. E., and Zamora, N. A. 

1990. Nyssa talamancana (Cornaceae) an addition to the rem- 
nant Laurasian Tertiary flora ofsouthern Central America. 
Brittonia, vol. 43, pp. 165-170. 

Hanson, C. B. 

1973. Geology and vertebrate faunas in the type area of the Сато 
Formation, Oregon. Geological Society of America, Cor- 
dilleran Section Abstracts with Programs, vol. 5, pp. 50. 

1991. Telataceras radinskyi, a new primitive rhinocerotid from 
the late Eocene Clarno Formation, Oregon, in Prothero, 
D. R. and Schoch, R. M. [eds.], The Evolution of Peris- 
sodactyls, pp. 379-398, Oxford Monographs on Geology 
and Geophysics. Oxford University Press, New York. 

Herendeen, P. S. 

1991. Lauraceous wood from the mid-Cretaceous Potomac Group 
of eastern North America: Paraphyllanthoxylon marylan- 
dense sp. nov. Review of Palaeobotany and Рајупојову, 
vol. 69, pp. 277-290. 

Herendeen, P. S. and Dilcher, D. L. 

1990. Fossil mimosoid legumes from the Eocene and Oligocene 
of southeastern North America. Review of Palaeobotany 
and Palynology, vol. 62, pp. 339-361. 

Hergert, H. L. 

1961. Plant fossils in the Clarno Formation, Oregon. Ore Bin, 

vol. 23, pp. 55-62. 
Hill, A. W. 

1933. The method of germination of seeds enclosed in a stony 
endocarp. Annals of Botany, vol. 47, pp. 873-887. 

1937. The method of germination of seeds enclosed in a stony 
endocarp. II. Annals of Botany (New Series), vol. 1, PP- 
239-256. 

Hollick, A. 

1936. The Tertiary floras of Alaska. U.S. Geological Survey Pro- 

fessional Paper 182, pp. 1-185. 
Holy, F. 

1975. Representatives of the family Mastixiaceae Calestani 1 905 
in the Bohemian Tertiary. Sbornik Närodniho Musea У 
Praze; Acta Musei Nationalis Pragae, vol. 31B, pp. 123- 
147, pl. 1—4. 


| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 125 


Howard, R. A. 
1942. Studies of the Icacinaceae IV: Consideration of the New 
World genera. Contributions ofthe Gray Herbarium, vol. 
142, pp. 1-60. 
Huzioka, K. and Takahasi, E. 
1970. The Eocene flora ofthe Ube Coal-field, southwest Honshu, 
Japan. Journal ofthe Mining College of Akita University, 
Ser. A, vol. 4, по. 5, pp. 1-88. 
Jähnichen, H. 
1976. Schisandraceae und Illiciaceae als holarktische und sub- 


tropische Florenelemente im mitteleuropäischen Alttertiär. - 


Abhandlungen des Zentralen Geologischen Instituts Pa- 
läontologische Abhandlungen, vol. 26, pp. 151-197, pl. 
8-18. 

Jain, В. К. 

1963. Studies in Musaceae— 1. Musa cardiosperma sp. nov., a 
fossil banana fruit from the Deccan Intertrappean Series, 
India. The Palaeobotanist, vol. 12, pp. 45-58, pl. 1-4. 

1965. Studies in Musaceae III. Fossil records of Musaceae and 
the Origin of Bananas. Proceedings of the Indian Academy 
of Science, vol. 61, pp. 170-179. 

Jones, J. Н. and Dilcher, D. L. 

1980. Investigations of angiosperms from the Eocene of North 
America: Rhamnus marginatus (Rhamnaceae) reexam- 
ined. American Journal of Botany, vol. 67, pp. 959-967. 

Kapil, R. N. and Jalan, S. 

1964. Schisandra Michaux—it embryology and systematic po- 

sition. Botaniska Notiser, vol. 117, pp. 285-306. 
Kaul, R. B. 

1988. Cupular structure in paleotropical Castanopsis (Fagaceae). 
Annals of the Missouri Botanical Garden, vol. 75, pp. 
1480-1498. 

1989. Fruit structure and ecology in paleotropical Lithocarpus 
(Fagaceae), in Crane, P. R. and Blackmore, S. [eds.]. Evo- 
lution, Systematics, and fossil history ofthe Hamamelidae, 
Volume 2: ‘Higher’ Hamamelidae. Systematics Associa- 
tion Special Volume no. 40B, pp. 67-86, Clarendon Press, 
Oxford, England. 

Kirchheimer, F. 

1936. Zur Kentniss der Friichte rezenter und fossiler Mastixioi- 
deen. Beihefte zum Botanischen Centralblatt, vol. 55, pp. 
275-300, pl. 5-8. 

1938. Beiträge zur näheren Kenntnis von Vitaceen-Samenfor- 
men tertiären Alters. Planta, vol. 28, pp. 582-598, pl. 5-8. 

1940. Die Mastixioideenflora der altertiàren Braunkohlenschi- 
chten von Wiesa bei Kamenz (Sachsen). 10 Jahresheft (1940) 
der Naturwissenschaftlichen Gesellshaft Isis in Kamenz, 
C.S. Krausche, Kamenz (Sachsen), 32 pp. 

1949. Die Symplocaceae der erdgeschichtlichen Vergangenheit. 
Palaeontographica, Abt. B, vol. 90, pp. 1-52, pl. 1-2. 

1957. Die Laubgewächse der Braunkohlenzeit. Veb Wilhelm 
Knapp Verlag, Halle (Salle), 783 pp. 

Knobloch, E. and Mai, D. H. 

1986. Monographie der Früchte und Samen in der Kreide von 
Mitteleuropa. Vydal Ustredni ústav geologický, vol. 47, 
рр. 219, 56 pl. 

Kovacs, E. 

1957. Dracontomelon minimum Reid-Chandler aus den Eozän- 
Kohlenschichten von Tatabanya. Földtany Közlöny; Bul- 
letin ofthe Hungarian Geological Society, vol. 87, pp. 215- 
217, 

Krause, J. 

1942. Staphyleaceae. Die Natürlichen Pflanzenfamilien, 2, vol. 

206, pp. 255-321. 


Krützsch, W. 

1976. Die Mikroflora der Braunkohle des Geiseltales, Teil IV: 
Die stratigraphische Stellung des Geiseltalprofils im Eozän 
und die sporenstratigraphische Untergliederung des mit- 
tleren Eozäns. Abhandlungen des Zentralen Geologischen 
Instituts Berlin vol. 26, pp. 47-92. 

1989. Paleogeography and historical phytogeography (paleocho- 
rology) in the Neophyticum. Plant Systematics and Evo- 
lution, vol. 162, pp. 5-61. 

Kvaéek, Z. 

1982. Tertiary taxads of NW Bohemia. Acta Universitatis Car- 

olinae — Geologica, Pokornyaa, vol. 4, pp. 471-491. 
Куасек, Z., Büzek, С., and S. R. Manchester. 

1991. Fossil fruits of Pteleaecarpum Weyland— Tiliaceous not 

sapindaceous. Botanical Gazette, vol. 152, pp. 522-523. 
Kvacek, Z. and Walther, H. 

1989. Paleobotanical studies in Fagaceae of the European Ter- 
tiary. Plant Systematics and Evolution, vol. 162, pp. 213- 
229. 

Lam, H. J. 

1931. Beiträge zur Morphologie der dreizähligen Burseraceae- 
Canarieae. Annales du jardin botanique de Buitenzorg, 
vol. 42, pp. 23-56, 3 pl. 

1932. Beiträge zur Morphologie der Burseraceae. Inbesondere der 
Canarieae. Annales du jardin botanique de Buitenzorg, 
vol. 42, pp. 97—226, 6 pl. 

Laurent, L. 

1912. Florefossile des schistes de Menat (Puy-de-Dóme). Annales 
du Musée d'Histoire Naturelle de Marseille, vol. 14, pp. 
1-246. 

Lesquereux, L. 

1878. Contributions to the flora of the Western Territories II. The 
Tertiary flora. Report of the U.S. Geological Survey of the 
Territories, vol. 7, pp. 1-366. 

Lockwood, E. H. and MacMillan, R. H. 

1978. Geometric symmetry. Cambridge University Press, Cam- 

bridge, England, 228 pp. 
Mabberley, D. J. 

1989. The plant-book. Cambridge University Press, Cambridge, 

England, 706 pp. 
MacGinitie, H. D. 

1969. The Eocene Green River flora of northwestern Colorado 
and northeastern Utah. University of California Publica- 
tions in Geological Sciences, vol. 83, 203 pp., 31 pl. 

1974. An early Middle Eocene flora from the Yellowstone-Ab- 
saroka Volcanic Province, northwestern Wind River Basin, 
Wyoming. University of California Publications in Geo- 
logical Sciences, vol. 108, 103 pp., 45 pl. 

Magallon-Puebla, S. and Cevallos-Ferriz, S. R. S. 

1992. Cedrelospermum (Ulmaceae) fruits from Upper Cenozoic 
Strata of Puebla, Mexico. American Journal of Botany, 
vol. 79(6), p. 103. Abstract. 

Mai, D. H. 

1970. Subtropische Elemente im europäischen Tertiär I. Die Gat- 
tungen Gironniera, Sarcococca, Illicium, Evodia, Ilex, 
Mastixia, Alangium, Symplocos und Rehderodendron. 
Paläontologische Abhandlungen Abt. В, vol. 3, pp. 441- 
503, pl. 58-69. 

1971. Uber fossile Lauraceae und Theaceae in Mitteleuropa. 
Feddes Repertorium, vol. 82, pp. 313-341. 

1975. Beitrage zur Bestimmung und Nomenklatur fossiler Mag- 
nolien. Feddes Repertorium, vol. 86, pp. 559-578. 

1976. Fossile Früchte und Samen aus dem Mitteleozän des Gei- 


126 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


seltales. Abhandlungen des Zentralen Geologischen Insti- 
tuts Berlin, vol. 26, pp. 93-149, pl. 1-7. 

1980. Zur Bedeutung von Relikten in der Florengeschichte. 100 
Jahre Arboretum (1879-1979), pp. 281-307. 

1984. Karpologische Untersuchungen der Steinkerne fossiler und 
rezenter Amygdalaceae (Rosales). Feddes Repertorium, vol. 
95, pp. 299-329, pl. 29-50. 

1985. Beiträge zur Geschichte einiger holziger Saxifragales-Gat- 
tungen. Gleditschia, vol. 13, pp. 75-88. 

1986. Über antillanische Symplocaceae. Feddes Repertorium, 
vol. 97, pp. 1-28. 

1987. Neue Früchte und Samen aus paläozänen Ablagerungen 
Mitteleuropas. Feddes Repertorium, vol. 98, pp. 197-229, 
pl. 12-21. 

1989. Development and regional differentiation ofthe European 
vegetation during the Tertiary. Plant Systematics and Evo- 
lution, vol. 162, pp. 79-91. 

Mai, D. H. and Walther, H. 

1978. Die Floren der Haselbacher Serie im Weisselster-Becken 
(Bezirk Leipzig, DDR). Abhandlungen der Staatliches Mu- 
seums für Mineralogie und Geologie, vol. 28, 200 pp., 50 
pl. 

1988. Die pliozänen Floren von Thüringen, Deutsche Demok- 
ratische Republik. Quartärpaläontologie, vol. 7, pp. 55- 
297, 41 pl. 

Manchester, S. R. 

1976. Description and interpretation of fossil woods, leaves and 
fruits from the Clarno Nut Beds, Wheeler County, Oregon. 
Oregon Museum of Science and Industry Technical Re- 
ports, vol. 4, pp. 65-102. 

1977. Wood of Tapirira (Anacardiaceae) from the Paleogene 
Clarno Formation of Oregon. Review of Palaeobotany and 
Palynology, vol. 23, pp. 119-127. 

1979. Triplochitioxylon (Sterculiaceae): A new genus of wood 
from the Eocene of Oregon and its bearing on xylem evo- 
lution in the extant genus Triplochiton. American Journal 
of Botany, vol. 66, pp. 699—708. 

1980. Chattawaya (Sterculiaceae): A new genus of wood from the 
Eocene of Oregon and its implications for xylem evolution 
of the extant genus Pterospermum. American Journal of 
Botany, vol. 67, pp. 59-67. 

1981. Fossil plants of the Eocene Clarno Nut Beds. Oregon Ge- 
ology, vol. 43, pp. 75-81. 

1983. Fossil wood of the Engelhardieae (Juglandaceae) from the 
Eocene: Engelhardioxylon gen. nov. Botanical Gazette, vol. 
144, pp. 157-163. 

1986. Vegetative and reproductive morphology of an extinct plane 
tree (Platanaceae) from the Eocene of western North Amer- 
ica. Botanical Gazette, vol. 147, pp. 200-226. 

1987a. The fossil history of the Juglandaceae. Monographs in sys- 
tematic botany from the Missouri Botanical Garden, vol. 
21, pp. 1-137. 

1987b. Extinct ulmaceous fruits from the Tertiary of Europe and 
western North America. Review of Palaeobotany and Pal- 
ynology, vol. 52, pp. 119-129. 

1988. Fruits and seeds of Tapiscia (Staphyleaceae) from the mid- 
dle Eocene of Oregon, USA. Tertiary Research, vol. 9, pp. 
59—66. 

1989a. Attached reproductive and vegetative remains of the extinct 
American- European genus Cedrelospermum (Ulmaceae) 
from the Early Tertiary of Utah and Colorado. American 
Journal of Botany, vol. 76, pp. 256-276. 

1989b. Early history of the Juglandaceae. Plant Systematics and 
Evolution, vol. 162, pp. 231-350. 


1989c. Systematics and fossil history of the Ulmaceae, in Crane, 
P. R. and Blackmore, S. [eds.]. Evolution, Systematics, and 
fossil history of the Hamamelidae, Volume 2: ‘Higher’ 
Hamamelidae. Systematics Association Special Volume 
no. 40B, pp. 221-252, Clarendon Press, Oxford, England. 

1990. Eocene to Oligocene floristic changes recorded in the Clar- 
no and John Day Formations, Oregon, USA, in E. Knob- 
loch, and Kvaéek, Z., [eds.], Symposium Proceedings, pa- 
leofloristic and paleoclimatic changes in the Cretaceous 
and Tertiary, pp. 183-187, Geological Survey Press, Prague, 
Czechoslovakia. 

1991. Cruciptera, a new juglandaceous winged fruit from the Eo- 
cene and Oligocene of western North America. Systematic 
Botany, vol. 16, pp. 715-725. 

Manchester, S. R. and Crane, P. R. 

1983. Attached leaves, inflorescences, and fruits of Fagopsis an 
extinct genus of fagaceous affinity from the Oligocene Flor- 
issant flora of Colorado, U.S.A. American Journal of Bot- 
any, vol. 70, pp. 1147-1164. 

Manchester, S. R. and Meyer, H. W. 

1987. Oligocene fossil plants from the John Day Formation, Fos- 

sil, Oregon. Oregon Geology, vol. 49, pp. 115-127. 
Manchester, S. R. and Miller, R. B. 

1978. Tile cells and their occurrence in malvalean fossil woods. 

IAWA Bulletin, vol. of 1978, pp. 23-28. 
Manchester, S. R. and Wheeler, E. A. 

1993. Extinct juglandaceous wood from the Eocene of Oregon 
and its implications for xylem evolution in the Juglanda- 
ceae. IAWA Journal, vol. 14, pp. 103-111. 

Manchester, S. R. and Zavada, M. S. 

1987. Lygodium foliage with intact sorophores from the Eocene 

of Wyoming. Botanical Gazette, vol. 148, pp. 392-399. 
Manchester, S. R. and Kress, W. J. 

1993. Fossil bananas (Musaceae): Ensete oregonense sp. NOV. 
from the Eocene of western North America and its phy- 
togeographic significance. American Journal of Botany, 
vol. 80, pp. 1264-1272. 

Mathewes, R. and Brooke, R. C. 

1971. Fossil Taxodiaceae and new angiosperm macrofossils from 

Quilchena, British Columbia. Syesis, vol. 4, pp. 209-216. 
Matthew, K. M. 

1976. Arevision of the genus Mastixia (Cornaceae). Blumea, vol. 

23, pp. 51-93. 
Mazer, 5. J. and Tiffney, B. Н. 

1982. Fruits of Wetherellia and Palaeowetherellia (? Euphorbi- 
aceae) from Eocene sediments in Virginia and Maryland. 
Brittonia, vol. 34, pp. 300-333. 

McClintock, E. 

1957. A monograph of the genus Hydrangea. Proceedings of the 
California Academy of Sciences, fourth series, vol. 29, pP- 
147-256. 

McGahan, М. У. 

1961. Studies on the seed of banana. I. Anatomy of the seed and 
embryo of Musa balbisiana. American Journal of Botany; 
vol. 48, pp. 230-238. 

McKee, T. M. 

1970. Preliminary report on fossil fruits and seeds from the Mam- 
mal Quarry of the Clarno Formation, Oregon. Ore Bin, 
vol. 32, pp. 117-132. 

McKenna, M. C. 

1975. Fossil mammals and early Eocene North Atlantic land 
continuity. Annals of the Missouri Botanical Garden, vol. 
62, pp. 335-353. 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 127 


Mellet, J. S. 

1969. 4 skull of Hemipsalidon (Mammalia Deltatheridia) from 
the Clarno Formation of Oregon. Novitates, American 
Museum of Natural History, no. 2387, pp. 1-19. 

Metcalfe, D. R. and Chalk, L. 

1950. Anatomy of the Dicotyledons. Clarendon Press, Oxford, 2 

vol., 1500 pp. 
Meyer, H. W. 

1992. Lapse rates and other variables applied to estimating pa- 
leoaltitudes from fossil floras. Palaeogeography, Palaeo- 
climatology, Palaeoecology, vol. 99, pp. 71-99. 

Mohana Rao, P. R. 

1974. Seed anatomy in some Hamamelidaceae and phylogeny. 

Phytomorphology, vol. 24, pp. 113-139. 
Morgan, D. R. and Soltis, D. E. 

1993. Phylogenetic relationships among members of Saxifraga- 
ceae sensu lato based on rbcL sequence data. Annals of 
the Missouri Botanical Garden, vol. 80, pp. 631-660. 

Muller, J. and Leenhouts, P. W. 

1976. A general survey of pollen types in Sapindaceae in relation 
to taxonomy, in Ferguson, I. К. and Muller, J. [eds.], The 
evolutionary significance of the exine, pp. 407-445. Aca- 
demic Press, London. 

Nilsen, T. H. and McKee, E. H. 

1979. Paleogene paleogeography of the western United States, in 
J. M. Armentrout, Cole, M. R., and TerBest, H., Jr. [eds.], 
Cenozoic paleogeography of the western United States. pp. 
257-276, Pacific Coast Paleogeography Symposium 3. Pa- 
cific Section, SEPM. 

Nixon, K. C. 

1989. Origins of Fagaceae, in Crane, P. R. and Blackmore, S. 
[eds.]. Evolution, Systematics, and fossil history of the 
Hamamelidae, Volume 2: ‘Higher’ Hamamelidae. Sys- 
tematics Association Special Volume no. 40B, pp. 23-44, 
Clarendon Press, Oxford, England. 

Noblett, J. B. 

1981. Subduction-related origin of the volcanic rocks of the Eo- 
cene Clarno Formation near Cherry Creek, Oregon. Or- 
egon Geology, vol. 43, pp. 91-99. 

Nooteboom, H. P. 

1975. Revision of the Symplocaceae of the Old World New Cal- 
edonia excepted. Universitair Pers Leiden, The Nether- 
lands, 335 pp. 

1985. Notes on Magnoliaceae with a revision of Pachylarnax and 
Elmerrillia and the Malesian species of Manglietia and 
Michelia. Blumea, vol. 31, pp. 65-121. 

Parrish, J. T. 

1987. Global palaeogeography and palaeoclimate of the Late 
Cretaceous and Early Tertiary, in E. M. Friis, Chaloner, 
W.G., and Crane, P. К. [eds.], The origins of angiosperms 
and their biological consequences, pp. 51—73. Cambridge 
University Press, Cambridge, England. 

Pigg, К. A. and Stockey, В. A. 

1991. Platanaceous plants from the Paleocene of Alberta, Can- 
ada. Review of Palaeobotany and Palynology, vol. 70, pp. 
125-146. 

Potbury, S. S. 

1935. The La Porte flora of Plumas County, California. Carnegie 
Institute of Washington Publication 465, pp. 29-82, pl. 
1019. 

Radlkofer, L. 

1890. Ueber die Gliederung der Familie der Sapindaceae. Sit- 

zungsberichte der mathemathematisch-physikalischen 


Classe de Bayerische Akademie Wissenschaften München, 
vol. 20, pp. 105-379. 
Reid, E. M. and Chandler, M. E. J. 

1933. The London Clay Flora. British Museum (Natural His- 

tory), London, England, vii + 561 pp., 33 pl. 
Retallack, G. 

1981. Preliminary observations on fossil soils in the Clarno For- 
mation (Eocene to early Oligocene) near Clarno, Oregon. 
Oregon Geology, vol. 43, pp. 147-150. 

1991. 4A field guide to mid-Tertiary paleosols and paleoclimatic 
changes in the high desert of central Oregon— Part 1. Or- 
egon Geology, vol. 55, pp. 51-59. 

Rogers, J. J. W. and Novitsky-Evans, J. M. 

1977. The Clarno Formation of central Oregon, U.S.A.—volca- 
nism on a thin continental margin. Earth and Planetary 
Science Letters, vol. 34, pp. 56-66. 

Rohrer, J. R., Robertson, K. R., and Phipps, J. B. 

1991. Variation in structure among fruits of Maloideae (Rosa- 
ceae). American Journal of Botany, vol. 78, pp. 1617- 
1635. 

Roth, J. L., Jr. and Dilcher, D. L. 

1979. Investigations of angiosperms from the Eocene of North 
America: Stipulate leaves of the Rubiaceae including a 
probable polyploid population. American Journal of Bot- 
any, vol. 66, pp. 1194—1207. 

Rüffle, L. 

1976. Myricaceae, Leguminosae, Icacinaceae, Sterculiaceae, 
Nymphaeaceae, Monocotyledones, Coniferae. Eozäne Flo- 
ren des Geiseltals. Abhandlungen des Zentralen Geolo- 
gischen Instituts Berlin, vol. 26, pp. 337—438. 

Saleeby, J. B. 

1983. Accretionary tectonics of the North American Cordillera. 
Annual Review of Earth and Planetary Science, vol. 15, 
pp. 45-73. 

Salisbury, E. J. 

1942. The reproductive capacity of plants. Studies in quantitative 

biology. G. Bell and Sons, London, England, 244 pp. 
Schaarschmidt, F. 

1988. Der Wald, fossile Pflanzen als Zeugen eines warmen Kli- 
ma, in S. Schaal, and Ziegler, W. [eds.] Messel—Ein 
Schaufenster in die Geschichte der Erde und des Lebens, 
pp. 29-52, W. Kramer, Frankfurt am Main, Germany. 

Schenk, E. T. and McMasters, J. H. 

1956. Procedure in Taxonomy. Stanford University Press, Stan- 

ford, California. 3rd ed., 119 pp. 
Schmid, R. 

1986. Oncornerian and other terminology of angiospermous and 
gymnospermous seed coats: Historical perspective and ter- 
minological recommendations. Taxon, vol. 35, pp. 476- 
491. 

Schumann, K. 

1897. Rubiaceae. in Engler and Prantl, Die natürlichen Pflan- 
zenfamilien. Teil 4, Abteilung 4, pp.*1-194, W. Engel- 
mann, Leipzig. 

Scott, A. and Collinson, М. Е. 

1983. Investigating fossil plant beds, part 1: the origin of fossil 
plants and their sediments. Geology Teaching, vol. 7, pp. 
114-122. 

Scott, R. A. 

1954. Fossil fruits and seeds from the Eocene Clarno Formation 
of Oregon. Palaeontographica, Abt. B, vol. 96, pp. 66-97, 
pl. 15-16. 

1956. Evolution of some endocarpal features in the tribe Tinos- 
poreae (Menispermaceae). Evolution, vol. 10, pp. 74-81. 


128 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Scott, R. A., Barghoorn, E. S., and Prakash, U. 

1962. Wood of Ginkgo in the Tertiary of western North America. 

American Journal of Botany, vol. 49, pp. 1095-1101. 
Scott, R. A. and Wheeler, E. A. 

1982. Fossil woods from the Eocene Clarno formation of Oregon. 

IAWA Bulletin, n.s., vol. 3, pp. 135-154. 
Sela, J. and Boyde, A. 

1977. Cyanide removal of gold from SEM specimens. Journal of 

Microscopy, vol. 3, pp. 229-231. 
Sellick, J. T. C. 

In Press. Phasmid eggs from the Eocene of Oregon. Palaeontology, 

vol. 36. 
Sleumer, H. 

1942. Icacinaceae. in Engler and Prantl, Die natürlichen Pflan- 
zenfamilien, 20b. Aufl. 2. Leipzig. 

1969. Materials toward the knowledge of the Icacinaceae of Asia, 
Malesia, Australia and the Pacific. Blumea, vol. 17, pp. 
181-264. 

1971. Icacinaceae. Flora Malesiana, ser. I, vol. 7, pp. 1-87. 

Smith, А. С. 

1947. The families Illiciaceae and Schisandraceae. Sargentia, vol. 
7, pp. 1-224. 

Standley, P. C. and Williams, L. O. 

1975. Flora of Guatemala. Rubiaceae. Fieldiana Botany, vol. 
124, pp. 1-274. 

Stirton, R. A. 

1944. A rhinoceros tooth from the Clarno Eocene of Oregon. 

Journal of Paleontology, vol. 18, pp. 265-267. 
Stockey, R. A. 

1984. Middle Eocene Pinus remains from British Columbia. Bo- 

tanical Gazette, vol. 74, pp. 1878-1887. 
Stockey, В. A. and Crane, Р. К. 

1983. In Situ Cercidiphyllum-like seedlings from the Paleocene 
of Alberta, Canada. American Journal of Botany, vol. 70, 
pp. 1564-1568. 

Sun, F. and Stockey, R. A. 

1992. A new betulaceous plant from the Paleocene of Alberta, 
Canada. International Journal of Plant Science, vol. 153, 
pp. 136-146. 

Swanson, D. A. and Robinson, P. T. 

1968. Base of the John Day Formation in and near the Horse 
Heaven mining district, north-central Oregon. U.S. Geo- 
logical Survey Professional Paper 600-D, pp. 154-161. 

Takaso, T. and Tobe, H. 

1990. Seed coat morphology and evolution in Celtidaceae and 
Ulmaceae (Urticales). Botanical Magazine, Tokyo, vol. 103, 
pp. 25-41. 

Takhtajan, A. 

1966. Major phytochoria of the late Cretaceous and the early 
Tertiary in USSR and adjacent countries. 13th Sir Charles 
Seward Memorial lecture Lucknow, India, 12 pp. 

1986. Floristic regions ofthe world. University of California Press, 
Berkeley, 522 pp. 

Tanai, T. 

1990. Euphorbiaceae and Icacinaceae from the Paleogene of 
Hokkaido, Japan. Bulletin of the National Science Mu- 
seum, series C (Geology and Paleontology), vol. 16, pp. 
91-118. 

Tanai, T. and Wolfe, J. A. 

1977. Revisions of Ulmus and Zelkova in the middle and late 
Tertiary of western North America. U.S. Geological Survey 
Professional Paper 1026, pp. 1-11. 

Taylor, D. W. 
1990. Paleobiogeographic relationships of angiosperms from the 


Cretaceous and early Tertiary ofthe North American area. 
Botanical Review, vol. 56, pp. 279-417. 
Teichman, I., van, and Robbertse, P. J. 

1986. Development and structure ofthe pericarp and seed of Rhus 
lancea L. fil. (Anacardiaceae), with taxonomic notes. Bo- 
tanical Journal of the Linnean Society, vol. 93, pp. 291- 
306. 

Thanikaimoni, G. 

1984. Ménispermacées: Palynologie et Systématique. Institut 
Francais de Pondichery Travaux de la Section Scientifique 
et Technique, Pondichery, India, vol. 18, 135 pp. 

1986. Evolution of Menispermaceae. Canadian Journal of Bot- 
any, vol. 64, pp. 3130-3133. 

Tiffney, B. H. 

1977. Fruits and seeds of the Brandon Lignite: Magnoliaceae. 
Botanical Journal of the Linnean Society, vol. 75, pp. 299- 
2 

1979. Fruits and seeds of the Brandon Lignite, III. Turpinia 
(Staphyleaceae). Brittonia, vol. 31, pp. 39-51. 

1981. Fruits and seeds of the Brandon Lignite, VI. Microdiptera 
(Lythraceae). Journal of the Arnold Arboretum, vol. 62, 
pp. 487-516. 

1984. Seed size, dispersal syndromes, and the rise of the angio- 
sperms: evidence and hypothesis. Annals of the Missouri 
Botanical Garden, vol. 71, pp. 551-576. 

1985a. Perspectives on the origin of the floristic similarity between 
eastern Asia and eastern North America. Journal of the 
Arnold Arboretum, vol. 66, pp. 73-94. 

1985b. The Eocene North Atlantic land bridge: its importance in 
Tertiary and modern phytogeography of the Northern 
Hemisphere. Journal of the Arnold Arboretum, vol. 66, 
pp. 243-273. 

1990. The collection and study of dispersed angiosperm fruits and 
seeds. Palaios, vol. 5, pp. 499—519. 

Tiffney, B. H. and Barghoorn, E. S. 

1976. Fruits and seeds of the Brandon Lignite: I. Vitaceae. Ке- 
view of Palaeobotany and Palynology, vol. 22, pp. 169- 
191. 

Traverse, А. 

1955. Pollen analysis of the Brandon Lignite of Vermont. U.S. 

Bureau of Mines Report of Investigations 5151, 107 pP- 
Upchurch, G. R., Jr., and Wolfe, J. A. 

1987. Mid-Cretaceous to Early Tertiary vegetation and climate: 
evidence from fossil leaves and woods, in E. M. Friis, Chal- 
oner, W. G., and Crane, P. R. [eds.], The origins of an- 
giosperms and their biological consequences, pp. 75-106. 
Cambridge University Press, Cambridge, England. 

Vance, J. А. 

1988. New fission track and K-Ar ages from the Clarno For- 
mation, Challis age volcanic rocks in north central Oregon. 
Geological Society of America, Rocky Mountain Section 
Abstracts with Programs, vol. 20, p. 473. 

Walker, G. W. 

1977. Geologic map of Oregon east of the 121st meridian. US. 
Geological Survey Miscellaneous Investigation Series Мар 
1-902, scale 1:500,000. 

Walker, G. W. and Robinson, P. I. 

1990. Paleocene (?) Eocene, and Oligocene (?) rocks of the Blue 
Mountain Region, in G.W. Walker, [ed.]. Geology of the 
Blue Mountains Region of Oregon, Idaho, and Washing- 
ton: Cenozoic geology of the Blue Mountains Region. PP- 
13-28. U.S. Geological Survey Professional Paper 1437. 

Wang, C.-W. 
1961. The forests of China with a survey of grassland and deser 1 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 129 


vegetation. Maria Moors Cabot Foundation publication 
no. 5, Cambridge, Massachusetts, 313 pp. 
Welzen, P. C., van 

1988. Key to the Malesian genera of Sapindaceae (based on veg- 
etative and fruit characteristics). Blumea, vol. 33, pp. 215— 
28 

Werft, H., van der 

1991. Akeytothe genera of Lauraceae inthe New World. Annals 

of the Missouri Botanical Garden, vol. 78, pp. 377-387. 
Wheeler, E. and Baas, P. 

1992. Fossil wood of the Leguminosae: a case study in xylem 
evolution and ecological anatomy, in Herendeen, Р. $. and 
Dilcher, D. L. [eds.], Advances in Legume systematics part 
4 the fossil record, pp. 281-302, Royal Botanic Gardens, 
Kew, England. 

Wilde, У. 

1989. Untersuchunge zur Systematik der Blattreste aus dem Mit- 
teleozän der Grube Messel bei Darmstadt (Hessen, Bun- 
desrepublik Deutschland). Courier Forschungsinstitut 
Senckenberg, vol. 115, pp. 1-213. 

Wing, S. L. 

1988. Eocene and Oligocene floras and vegetation of the Rocky 
Mountains. Annals of the Missouri Botanical Garden, vol. 
74, pp. 748-784. 

Wing, S. L. and Hickey, L. Н. 

1984. The Platycarya perplex and the evolution of the Juglan- 

daceae. American Journal of Botany, vol. 71, pp. 388- 


411. 
Wolfe, J. A. 

1968. Paleogene biostratigraphy of nonmarine rocks in King 
County, Washington. U.S. Geological Survey Professional 
Paper 571, pp. 1-33. 

1971. Tertiary climatic fluctuations and methods of analysis of 
Tertiary floras. Palaeogeography, Palaeoclimatology, 
Palaeoecology, vol. 9, pp. 27—57. 

1972. An interpretation of Alaskan Tertiary floras in Floristics 
and Paleofloristics of Asia and eastern North America, in 
Graham, A. [ed.], Floristics and Paleofloristics of Asia and 
eastern North America, pp. 201-233, Elsevier, Amster- 
dam. 

1975. Some aspects of plant geography of the Northern Hemi- 
sphere during the Late Cretaceous and Tertiary. Annals of 
the Missouri Botanical Garden, vol. 62, pp. 264-279. 

1977. Paleogene floras from the Gulf of Alaska Region. U.S. 
Geological Survey Professional Paper 997, pp. 1-107. 

1979. Temperature parameters of humid to mesic forests of east- 


ern Asia and relation to forests of other regions in the 
Northern Hemisphere. U.S. Geological Survey Profession- 
al Paper 1106, pp. 1-37. 

1981. A chronological framework for Cenozoic megafossil floras 
of northwestern North America and its relation to marine 
geochronology. Geological Society of America Special Pa- 
per, vol. 184, pp. 39-47. 

1985. Distribution of major vegetational types during the Terti- 
ary, in The carbon cycle and atmospheric CO,: Natural 
variation, Archaean to Present, Geophysical Monograph 
32, pp. 357-375. 

Wolfe, J. A. and Hopkins, D. M. 

1967. Climatic changes recorded by Tertiary land floras in North- 
western North America, in K. Hatai, [ed.], Tertiary Cor- 
relations and Climatic Changes in the Pacific, 11th Pacific 
Science Congress of the Pacific Science Association, Tokyo, 
vol. 25, pp. 67-76. 

Wolfe, J. A. and Poore, R. Z. 

1982. Tertiary marine and nonmarine climatic trends, in W. 
Berger, and Crowell, J.C. [eds.]. Climate in Earth history, 
pp. 154-158, National Academy of Science, Studies in 
Geophysics, Washington, D.C. 

Wolfe, J.A. and Tanai, T. 

1987. Systematics, phylogeny, and distribution of Acer (maples) 
in the Cenozoic of western North America. Journal of the 
Faculty of Science, Hokkaido University, vol. 22, pp. 1- 
246. 

Wolfe, J. A. and Upchurch, С. A., Jr. 

1986. Vegetation, climatic, and floral changes at the Cretaceous- 

Tertiary boundary. Nature, vol. 324, pp. 148-151. 
Wolfe, J. A. and Wehr, W. 

1987. Middle Eocene dicotyledonous plants from Republic, 
Northeastern Washington. U.S. Geological Survey Bulle- 
tin 1597, pp. 1-25. 

Wu T.-L. 

1981. Musaceae, Zingiberaceae, Cannaceae, Marantaceae, Bur- 
manniaceae. Flora Reipublicae Popularis Sinicae, vol. 16 
(2) Science Press, Beijing, pp. 1-189. 

Xiang, Qiu-Yun, Soltis, D. E., Morgan, D. R., and Soltis, P. S. 

1993. Phylogenetic relationships of Cornus L. sensu lato and pu- 
tative relatives inferred from rbcL sequence data. Annals 
of the Missouri Botanical Garden, vol. 80, pp. 723-734. 

Zona, S. 

1990. A monograph of Sabal (Arecaceae: Coryphoideae). Aliso, 

vol. 12, pp. 583-666. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 1* 


Figure Page 
1. Equistetum clarnoi Brown strobilus showing hexagonal sporangiophores. OMSI-Pb 582. RL, х3. ............................ 15 
ТОК пие bc deca n evt UH Od ee кышлык ee NE EO 30 
3. Compressed pollen cone of Pinus sp. from West Branch Creek locality 230. Note bulging basal scales for comparison with figs. 6-8. 

EA N ee N ША ЗЫ! М N ДИВИ earn СГ, 
4,5. Seeds 0D 725 SD. Кой Claro West Branch Creek locality 2296. ШЕ 9959 and 9960, XT. opiso eors e eee e E e a a 30 
TESTEN UE N НС ОРЕ RE ревија SERIE cae 30 


6. Longitudinally fractured specimen, showing basal portion of the cone with bulging scales. RL, х5. 

7. Same specimen, sectioned longitudinally near the periphery, showing helical arrangement of sporophylis. RL, х5. 
8. Same, sectioned longitudinally showing thick central axis. RL, х5. 

9. Longitudinal thin section showing pollen sacs on the abaxial side of the microsporophylls. RL, x 20. 


* For explanation of abbreviations used in Plate captions, see p. 30. 


PLATE 1 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PLATE 2 


a a> 
EA 


em?) 
RR 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 131 


EXPLANATION OF PLATE 2 


Figure Page 
EHER АЛ NA Ар ЛОМ II RCRUM I dL DIAM ee 31 
1. Seed, lateral view showing pointed арех апа truncate base. Holotype, USNM 355474. PC, х6. 
2. Same, face view, showing smooth surface, rounded-obtuse apex, truncate base. PC, x6. 
3, 4. Same, stereo-pair, apical view, showing dorsiventral keel. PC, x 6.4. 
5. Seed, lateral view. USNM 355406. PC, x 6.4. 
6. Same, apical view, pointed micropylar end. PC, x 6.4. 
7, 8. Same, stereo-pair, basal view, showing elliptical truncation with pair of vascular bundle scars. PC, x 6.4. 
9. Transverse section near base of the holotype, figs. 1-4, showing the two vascular traces (arrows). USNM 355474. RL, immersed 
in water, x20. 
10. Same, transverse section midway between base and apex, showing the lensoidal outline and uniform thickness of the seed 
coat. USNM 355474. RL, x10. 
11. Detail from fig. 10, showing isodiametric cells composing the seed coat. RL, х 50. 
= CeO Tren ac ООЛУ SD EDO PO tos ten И ЫЕ AP И MM кгс р C ee, 31 
12. Seed in face view, showing ovate outline and smooth surface. Holotype, UF 6510. 
13. Same, lateral view, showing apical keel and prominent vascular pore (arrow). 
14. Another example in face view. Lateral swellings of the margin correspond to the position of the two vascular pores. USNM 
422380. 
15. Same specimen as in figs. 12, 13, apical view, showing elliptical outline and median keel. 
16. Same specimen as in fig. 14, apical view, showing elliptical outline. 


132 


Figure 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 3 


lee: DIODOS LO EN OTE CS: Sp TONE Wate ne en a ee ae 


152. 


And ш 


Stereo-pair, specimen with aril preserved in the upper half with wrinkled, pitted surface. Seed cast exposed in lower half. 
USNM 355569. x5. 


. Same, basal view, showing thickness of the aril, which is partially broken away revealing smooth seed cast within. х5. 

. Same, apical view, showing micropylar aperture and wrinkly outer surface of the aril. х5. 

. Internal mold of seed coat, face view, showing wide-elliptical outline and faint longitudinal striations. UF 8541. PC, х5. 
. Seed in lateral view showing longitudinal striations of the aril, rounded base, dorsiventral keel extending over the apex and 


vascular pore (arrow). Holotype, UF 8542. PC, х5. 


. Same, apical view, showing elliptical outline, dorsiventral keel and central micropyle. PC, х 5. 

. Same, basal view, showing smooth surface and pair of pores representing vascular scars (arrows). PC, х5. 

. Seed in face view. UF 8529. PC, x4.5. 

. Same, lateral view. PC, х 4.5. 

. Same, stereo-pair, basal view, showing the smooth contours and the pair of lateral pores. PC, x4.5. 

. Median longitudinal section in the dorsiventral plane showing seed coat and the two lateral pores (arrows). UF 8513. Etched, 


КЗ. 


. Median longitudinal section in the dorsiventral plane of the specimen in fig. 5, showing elongate embryo. Etched, RL, х4.7. 
. Same, enlarged, showing folds of the embryo and speckled appearance of surrounding tissue. Etched, RL, x 10. 
. Transverse section of seed showing circular outline of the embryo in the center and partially intact aril surrounding the seed 


coat. UF 8514. RL, x5. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 3 


PLATE 4 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 133 


| EXPLANATION OF PLATE 4 


1-4. Actinidia AAO eat оду MENS Mec EN M C EM ee ra do ce а E 33 
1. Seed cast, lateral view, showing reticulate surface, bluntly pointed micropyle (m) and adjacent hilum (h). Holotype, UF 
6292. SEM, x13. 
2. Same, hilar end, showing lensoidal outline of seed. SEM, x18. 
3. Seed cast in matrix. UF 6498. SEM, x20. 
4. Same, enlarged to show space between outer and inner molds of the seed coat. SEM, x 50. 
5-14. ATA аер E A ee N AR RE 34 
5. Endocarp cast, face view, showing ovate outline. UF 6508. PC, x5. 
6. Same, lateral view, showing longitudinal septal groove and left carpel slightly larger than right carpel. PC, x5. 
7. Endocarp cast, face view, showing smaller carpel (front) and larger carpel (behind). USNM 422393. PC, x5. 
8. Same, lateral view, showing sculpture of septal wall and one carpel larger than the other. PC, x5. 
9. Locule cast, dorsal view, showing elliptical outline and smooth surface. USNM 424646. PC, x5. 
0. Same, lateral view, showing adjoining locule casts of a fruit with the endocarp wall eroded away, revealing the median 
septum with arms of the septal canal system emerging as a median row of circular protruberances. PC, х5. 
11, 12. Stereo-pair of a permineralized endocarp, face view, partially broken, revealing one of the locule casts. Holotype, USNM 
424646. x5. 
13. Same, transverse section, showing relatively thin wall, median septum perforated by canals, and two locules with remains 
of the seed coat in each. RL, x7. 
15 14. Isolated silica cast of the septal canal system. UF 9479. PC, x5. 
> 16. ОАА о) Nov: Holotype USNM ОООО ul s а eee ee, 35 
15. Permineralized endocarp, lateral view, showing greater thickness relative to figs. 6, 8. Arms of the septal canal system 
emerging as a median row of circular protruberances. PC, х5. 
16. Transverse section, showing thick lateral walls and locules virtually equivalent in size. X 7. 


134 


Figure 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 5 


1- 7 PERO PA С ИПИ ШУ eid et Chandler comb DOV ee: 


ы 


За 


> 


Stereo-pair, apical view of fruit, showing rounded pentagonal outline with five elliptical germination valves. USNM 422390. 
РС, 705. 
Same specimen, basal view, showing small central scar and relatively smooth surface. PC, x 5.5. 


. Same, lateral view, showing germination valves in apical half. PC, х5.5. 
. Isolated locule cast, lateral view, showing oblique apical truncation corresponding to the germination valve of the fruit. UF 


6664. PC, x5. 


. Same, dorsal view, showing median ridge along the apical truncation. PC, x5. 
. Successive transverse sections from apex to base of the specimen in figs. 1—4. RL, хб. 


7. Section near the apex, above the level of locules and central vascular strands. 

8. Section showing the central vascular bundles and five locules, each with a pair of shutter-like germination valves 
(arrows). 

9. Nearly equatorial section, showing interlocular lacunae. 

10. Equatorial section showing thick outer wall and five locules alternating with five lacunae. Seeds with embryos are 
preserved in two of the locules. 

11. More basal section, below the interlocular lacunae. 

12. Basalmost section. 


. Transverse section of a fruit in which one of the five germination valves has opened, admitting sediment to the locule. UF 


5708. RL, x6. 


. Apical view, specimen from the London Clay flora for comparison with the Clarno fossils. BM v30096. RL, х5.5. 
. Apical view of a rare tetracarpellate specimen. USNM 424851. PC, x5.5. 

. Same, lateral view. PC, x5.5. 

. Same, sectioned longitudinally. RL, x6. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 5 


PLATE 6 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 135 


EXPLANATION OF PLATE 6 


Figure Page 
i-e he NR eL EIN ишт d оли t cues 36 
1, 2. Stereo-pair, face view, specimen with longitudinally ribbed meso- and exocarp partially intact. Smooth surface of endocarp 
exposed at the upper right. Holotype, USNM 355036. RL, x7. 


3. Same, basal view, showing narrow outline and radiating ribs. RL, x 7. 

4. Endocarp, face view, showing smooth surface and rounded contours; diagonally broken at upper left. UF 9418. PC, x7. 

5. Same, oblique lateral view, with broken edge showing thick endocarp wall. SEM, x 11.5. 

6. Same, enlarged to show circular funicular scar and the layer of anticlinally elongate columnar cells in the endocarp wall. 
SEM, x25. 

7. Same, more greatly enlarged, showing surface cells of the endocarp arranged in rows radiating from the funicular depression. 
SEM, x40. 

8. Transverse section showing prominent layer of columnar cells in the endocarp. UF 9420. RL, x50. 

210: PROSA ОТТОНА СТА ЕО Е РСЗ E ee, TRU оу су л M 37 


9. Endosperm cast, face view, showing ellipsoidal outline, transverse ruminations and the encircling raphe band. UF 6574. 
10. Same, lateral view, showing longitudinal raphe band and transverse ruminations. 
11. Same, apical view. 
12. Smaller specimen, face view, showing transverse ruminations. UF 5233. 
13. Same, lateral view, showing raphe band. 
14. Larger specimen, face view, showing raphe band at the margin, transverse ruminations, and radially aligned ruminations 
near the base. UF 5234. 
15. Same, lateral view, showing raphe band partially broken away revealing vascular strand. 
18. EINONUSDERMIUNT DONESIL Spi NOV. AO lO pen OS NIVINS S897 3 RAS 38 
16, 17. Stereo-pair, endosperm cast, face view, with adhering remains of the seed coat with fibers arranged in a fingerprint pattern. 
18. Same, lateral view, showing narrow thickness. 


16- 


136 


Figure 
БЕЗ 


3,4. 


13—15. 


16—18. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 7 


ТОЛОО murni rotundum вре NOV. Holotype USNM 435099, BE x4, ee 38 
1. Face view showing wide elliptical outline and radial arrangement of ruminations. 
2. Same, lateral view showing seed thickness and course of the raphe. 
ПИ OIN ОЛО ONERI E S LIS Deal Vee DE соле ntis ig E н Be у а а 38 
3. Compressed fruit showing pedicel, D-shaped mericarps, epigynous perianth (arrows) and persistent recurved styles. Holotype, 
OMSI Pb810. 
4. Specimen with larger mericarps and persistent styles. OMSI Pb1779. 


. Extant Pseudopanax davidii (Franch.) W.R. Philipson, fruit for comparison with figs. 3 and 4. Western Hupeh, China. E.H. Wilson 


ПИ ОД RAS АЕ О ОИЕ р ара 38 


ООС ОСОО С СЕТОТ Е о ао OL nM TT 39 


6. Chalcedony nut cast, apical view, showing convergence of longitudinal striations toward the stylar projection. UF 8494. РС, 
за 25; 
7. Same, База! view, showing outer сисшаг scar of involucre attachment and an inner, more jagged circular scar where уазсшаг 
strands originating in the involucre passed into the base of the nut. PC, x 1.25. 
8. Same, lateral view, showing smooth rounded outline, longitudinal striations, basal involucre scar and apical stylar projection. 
PECX рада 
9. Nut cast, lateral view, showing longitudinal striations апа База! involucre scar. Holotype, UF 8497. PC, x 1.25. 
10. Same, basal view, showing jagged-margined circular area where the vascular strands passed into the involucre. PC, x 1.25. 
11. Siltstone mold from holotype specimen in figs. 9 and 10, showing prominant longitudinal ribs of vascular strands representing 
the impression of the outermost fruit layer. RL, x 1.25. 
12. Fractured nut with cast of the seed inside, basal view showing shriveled globose seed and circular hilum (arrow) within the 
fractured nut wall. UF 8498. PC, x 1.25. 
Extant Corylus sp. nut with involucre removed for comparison with Coryloides. UF mod. ref. coll. ........................ 39 
13. Lateral view showing globose form, smooth surface with merdional ribs, and prominent basal involucre scar for comparison 
with figs. 8, 9. RL, x 1.25. 
14. Basal view, showing irregular, more or less circular involucre scar with a narrow, zig-zag border for comparison with figs. 7, 
10. RL, x1.25. 
15. Transverse section showing wall of nut containing numerous large vascular bundles. RL, x2. 
СООТ ТОСО SR INE cd I east Se NE ca eene M Co A be кк MO dt а RERO 39 
16. Partially compressed specimen similar to that of fig. 11, showing longitudinal ribs. UF 9898. RL, x1.25. 
17. Same, transversely sectioned, showing prominence of the longitudinal ribs. Contrast with fig. 15. RL, x2. 
18. Same, transverse thin section, showing vascular bundles within the ribs. RL, х 5. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 7 


PLATE 8 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 137 


EXPLANATION OF PLATE 8 


Figure Page 
IB. КО ОООО Пату реВ CSD ОМ. SENE NT e UO па Тул а съ И D uec ke, а TM eg 40 
1. Locule cast, face view, showing median longitudinal groove and pointed apex. Holotype, USNM 435077. SEM, x 23. 
2. Another specimen, somewhat wider, face view, showing faintly cordate base. USNM 435076. SEM, x 21. 
3. Same as fig. 1, lateral view, showing narrow profile, pointed apex and rounded base. SEM, x 25. 
4. Apical view. USNM 424750. SEM, x21. 
5. Basal view. USNM 424076. SEM, x21. 
17. CUR DU TINO OSOHOHSISISP HOY Е ап Eo O RAT NT 40 
6. Endocarp, ventral view, showing ovoid outline and transverse placental slit. USNM 424887. PC, x5. 
7. Same, lateral view, showing rounded dorsal margin and ventral notch corresponding to placenta. PC, х 5. 
8, 9. Stereo-pair of endocarp, face view, showing transverse placenta and median ventral angle. Holotype, USNM 424885. PC, 
хе 
10. Same, apical view, showing triangular outline of the dorsal and ventral faces and the placenta slit seen on the ventral faces. 
Реж со 
11. Endocarp showing ventral placenta and ovoid outline. USNM 434955. РЄ; X. 
12. Same, dorsal view, with much of the endocarp wall fractured away, exposing the locule cast. PC, x5. 
13. Chalcedony fruit cast, lateral view, broken apically, with veins of exocarp preserved. USNM 355491. PC, x5. 
14. Same, basal view, showing angle of ventral faces and convex-rounded outline of the dorsal face. PC, x5. 
15. Median longitudinal section showing thickness of endocarp wall. UF 6648. RL, x7. 
16. Locule cast, ventral view showing prominent placental cavity. UF 6663. x7. 
17. Same, apical view. х7. 
18, 19. ESOO ED UNAS SIR ee eg ee ee ES оа И 41 
18. Endocarp, ventral view, showing median ventral angle and transverse placental slit. UF 6655. SEM, х 17. 
19. Same, lateral view. SEM, x 17. 
20. Betulaceous fruit similar to Palaeocarpinus with spiny bract arising from small nutlet similar in size to the locule casts of 
Таа West Branch теск оса 230 ЕЮ 20 en 40 


бе 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 9 


Figure Page 
О SIS ILO АЕ o кд E Corr c NP е M EE ГОИ АО T Es 
1. Endocarp, lateral view showing elliptical outline. Holotype, USNM 422378. PC, х7.5. 
3. Same, stereo-pair, rotated 90°, showing longiiudianal vascular grooves. PC, x 7.5. 
4. Shorter, wider endocarp, lateral view. USNM 424702. PC, x 7.5. 
5. Basal view of holotype as in figs. 1-3, showing circular outline and prominent vascular grooves. PC, x 7.5. 
6. Basal view of specimen in fig. 4. PC, x 7.5. 
7. Transverse section of specimen in figs. 1—3, and 5, showing two locules. Note absence of a median vascular bundle in the 
septum. RL, x10. 
8. Transverse section of specimen in figs. 4, 6. RL, x10. 
9-12. Trilocular specimen. USNM 355397. 
9. Lateral view, with pericarp partially broken away revealing shiny chalcedony locule cast. PC, x 7.5. 
10, 11. Stereo-pair, showing subglobose shape, locule cast seen in dorsal view. PC, x7.5. 
12. Transverse section showing the septum and three locules. RL, x8. 
13. Transverse section of the holotype, enlarged from in fig. 7, showing seed shriveled inside the locule, and vascular bundle 
distal from the septum (arrow). RL, x20. 
14. Same specimen, transversely sectioned closer to the apex, showing single layer of cuboidal cells surrounding the locules and 
forming zones of weakness along the planes of dehiscence (arrows). RL, x30. 
15. Peel from specimen in fig. 8, showing thick layer of isodiametric sclereids comprising the endocarp. TL, X 60. 


PLATE 9 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 10 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 139 


EXPLANATION OF PLATE 10 


Figure Page 
1-3. Mastixia SPIEL a Er HS ЖЫЕН. O ee M I че 42 
l. Transverse section of an endocarp with adhering matrix, showing smooth outer wall and outline of seed coat within the 
locule. UF 6502. Etched, RL, x 6.5. 
2. Transverse section of an endocarp in matrix showing typical thick dorsal infold and lines of dehiscence. UF 6303. RL, x 6.5. 
3. Transverse peel of endocarp wall, showing inner layer of horizontally aligned fibers, and an outer layer of obliquely aligned 
fibers. UF 6502. TL, x30. 
(ОООО ДО OI CEOE AS O S S ОЕ ll ә о? 43 
4. Transversely fractured fruit in matrix, showing prominently sculptured surface of endocarp and relatively narrow dorsal 
infold. UF 9207. RL, x3. 
5. Transverse section of fruit in matrix, with the surface sculpture worn by erosion, showing narrow dorsal infold, and a pair 
2 of lacunar canals adjacent to the lines of dehiscence. UF 9269. RL, х 5.5. 
6. Same, detail of wall, with lacunar canal. RL, x 15. 
7. Same, showing swirling tracts of fibers. RL, x 30. 
8. Latex cast prepared from the sedimentary mold of an endocarp, showing rough external sculpture and ovoid shape. UF 
9206. x 2.5. 
9-11. Fossil boat-shaped locule cast recovered from the mold of the specimen in fig. 8. x 3.5. 
9. Lateral view showing smooth surface, pointed apex, rounded base. 
10. Ventral view showing the convex surface. 
11. Dorsal view showing the median groove. 
gs ООО ter evi SALES: ONE oce A эу... су хык хыз ee ee. 44 
12. Fruit in matrix, broken and revealing elongate, boat-shaped locule cast. UCMP 10705. RL, х2.5. 
13. Transverse section of the same specimen showing thick wall including epicarp with smooth surface. Note seed within the 
locule and line of the dehiscence plane within the endocarp (arrow). RL, х3. 
14. Fruit in transverse section showing slight separation of epicarp from the endocarp, particularly apparent at lower left. 
Holotype, UF 8740. RL, х3. 
15. Same, detail showing swirling tracts of fibers in the endocarp and anticlinally oriented fibers of the epicarp. Note secretory 
canals in outer layer. RL, x15. 


чс 


I» 


140 


Figure 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 11 
Page 


ТЕТО Dnert Cand Reid ce Chandler s cue пне от оо аа 44 


L; 


л & C PN 


10. 


EIN E MALO aC OLE COUNT OVen Е МУ; ee Кы оек ЛИ TEE 


И 


127 
13-15. 


Complete endocarp in matrix, longitudinal view. UF 8731. x2. 


. Broken endocarp with the two locule casts protruding, longitudinal view. UF 8734. x2. 

. Apical view of the same specimen. х 2. 

. Etched transverse section of the endocarp from fig. 1, with one of the locules abortive. x2. 

. Isolated locule cast, ventral view, showing elongate, convex ellipsoidal outline and the median longitudinal raphe. UF 


8738. x2.5. 


. Same, dorsal view, showing pair of grooves corresponding to infolds of the endocarp. x2.5. 
. Transverse section of a typical endocarp showing thick wall, two fully developed w-shaped locules, and faint lines delimiting 


the germination valves (arrows). UF 8730. x2.5. 


. Transverse section of a small fruit with one fully developed locule. Note the epicarp preserved as a dark layer surrounding 


the endocarp; dorsal infolds (arrows) can be seen corresponding to an abortive locule, as well as those corresponding to 
the fully developed locule. USNM 424875. x2.5. 


. Acetate peel from fig. 8, magnified to show fibers making up the endocarp and remains of a seed within the underdeveloped 


locule. USNM 424875. TL, x30. 

Acetate peel from specimen in fig. 8, showing endocarp made up of variously oriented fibers, intruded and surrounded by 
the epicarp of thin-walled cells. Secretory canals present in the epicarp (arrows). TL, x30. , 
4 
Complete endocarp in matrix, longitudinal view showing ellipsoidal outline and equatorial plane of transverse section in 
fig. 16. UF 6842. x1.8. 

Abraded endocarp showing elliptical outline of germination valve. USNM 40543. x2.5. 

Isolated locule cast. UF 6838. x2.5. 

13. Ventral surface, showing obovate outline. 

14. Dorsal surface. 

15. Lateral view, showing curvature, ventral surface to the left. 


. Transverse section of fig. 11, showing three c-shaped locules, thick wall and undulate margins of the endocarp. х 2.0. 

. Apical view of fig. 12, showing the outline of 3 protruding locules casts delimiting three germination valves. x2.5. 

. Transverse section of a complete endocarp preserved in matrix. UF 6837. RL, x2.5. 

. Anatomical detail of endocarp from section in fig. 18, showing masses of fibers oriented in different directions. TL, х 50. 


PLATE 11 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 12 


6-11. 


12-15. 


16-18, 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 141 


EXPLANATION OF PLATE 12 


Page 
ENA SSCUMSGOELLIES[) поо d M UAM M Иа 46 
1. Chalcedony endocarp cast, lateral view, with one of the germination valves removed. Holotype, USNM 354588. PC, х4.5. 
2. Same, apical view, showing the three germination valves, the one on the left broken away. PC, x4.5. 


ISSUE ВЕ е RE Cont dtes ааа т Нети 46 


3. Large unilocular endocarp, lateral view, showing rounded apex and longitudinal surface grooves. UF 9757. x 1.5. 
4. Same, sectioned transversely, showing thick endocarp wall and single locule. RL, x 1.5. 
5. Same, detail of endocarp wall showing swirling tracts of fibers. RL, x30. 
EASTUNODSTSCRED ENL:SDANON BIOS DC ЕВ ОО seach es I c ыза ее авна ЖО. 46 
6. Complete fruit including cupule, viewed basally, showing circular outline. x 1.25. 
7. Same, viewed laterally, with portions of wall broken away revealing locule cast which is pointed apically and truncate basally. 
EIE 
8. Same, basal view, showing undulating circular outline of locule cast. x 1.25. 
9. Transverse section of the wall from fig. 6, showing dark endocarp layer surrounded by the adnate cupule. RL, x2. 
10. Detail of tranverse section, showing relatively dense tissue of the nutshell (lower half of photo), and less compact tissue of the 
cupule (upper half of photo). Note islands of concentrically arranged fibers in the nutshell. RL, x 12.5. 
11. Same, detail of nutshell, showing concentrically arranged fibers. TL, x75. 
Extant Castanopsis piriformis Hick. et A. Camus. Vietnam. M. Poilane 30313 (US 2493106). ............................. 47 
12. Apical view, showing relatively smooth circular outline, for comparison with fig. 6. x 1.25. 
13. Lateral view, showing saw cut for transverse section in fig. 14. x 1.25. 
14. Transverse section of the wall, showing nut surrounded by adnate cupule; cf. fig. 9. RL, x2. 
15. Same, thin section of the nutshell, showing islands of concentrically arranged fibers and an adjacent duct with dark contents; 
chang Е хто. 
ONErCUSSDALEOGALDA: Manchester К о в н 47 
16. Acorn, lateral view with nut and abraded cupule. Holotype, ОМІ Pb176. х 1.25. 
17. Same, apical view showing circular outline of the nut. x 1.25. 
18. Isolated cupule in matrix, lateral view, showing concentric transverse ribs. USNM 414508. x2. 


142 


Figure 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 13 


¡ELO SON TOP SUPR ОВО ОР ИО МУ O aa И, Е ОУ RN Inn 
1. A complete fruit viewed laterally showing the smooth surface, more or less rounded base and pointed apex. UF 6471. x4.2. 


- A similar fruit with some of the outer layer flaked away, revealing median longitudinal placental band. UF 6474. PC, х4. 
. A fruit with some of the wall removed in the apical half, exposing two rows of seeds attached to adjacent longitudinal placental 


bands. UF 6477. x4. 


. Stereo-pair of a specimen with large portion of outer wall broken away revealing numerous seeds. Holotype, UF 5236. x4. 
. Base of the specimen shown in fig. 2, showing four-fold symmetry, and four circular apertures corresponding to the placental 


bundles. PC, x5. 


. Specimen from fig. 1, sectioned transversely, showing the medial position of the placenta on each of the four valves of the 


fruit (arrows). Some of the seeds are preserved showing thin seed coat. Etched, RL, x8. 


. Transversely sectioned fruit in siltstone matrix, showing four parietal placentae with attached seeds. USNM 354008. RL, x8. 
. Same, enlarged more, showing arrangement of seeds around the placenta and pair of vascular bundles within the placenta. 


RE, x25. 


. Detail from fig. 5, showing ovoid seeds in attachment to the placenta. Subsequent transverse sectioning revealed that the row 


of seeds on the left are borne on the placenta to the left, while the row on the right corresponds to another placenta. SEM, 
x 8.5. 


. Longitudinally fractured fruit, showing dense mass of tortuous, more or less longitudinal, fibers and the impression of several 


seeds. UF 6575. SEM, x20. 


. Same, showing a seed fractured along the inside of the seed coat showing horizontally elongate hexagonal cells surrounded by 


fibers of the pericarp. SEM, x155. 


. Outer surface of a seed surrounded by fibers of the pericarp. SEM, x85. 

. Finer detail of same, showing the rectangular outline of seed coat cells. SEM, x400. 

. Detail from fig. 7, showing two seeds (arrows). Seed on right shows rectangular cells of seed coat. RL, x 120. 
. Seed from section in fig. 7, showing peripheral trichomes and shriveled inner layer of seed coat. RL, х 140. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 13 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 14 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 143 


EXPLANATION OF PLATE 14 


Figure Page 
1-13. Fortunearites endressii Пу Де ода по D eoque ce EC c NERIS Sen LEM IE cc cQ а 49 
1-5. Permineralized infructescence. Holotype, OMSI Pb156. (Counterpart of USNM 354581). 
1. Infructescence fractured longitudinally. OMSI Pb156. RL, х 1.5. 
2. Counterpart, showing stout axis and longitudinally fractured bilocular fruits. USNM 354581. RL, x2. 
3. Tangential section of the axis in fig. 2 showing the bilocular capsules in transverse view. RL, x3. 
4. Same, showing thin septum between locules and thick endocarp wall. RL, x 5. 
5. Transverse section of the axis in fig. 2, showing radiating fruits. RL, x2.5. 
6. Paired locule casts from a single fruit, face view. UF 9258. PC, x6. 
7. Same, apical view of the locule casts, cf. fig 4. PC, x6. 
8. Isolated locule cast removed from the infructescence in fig. 2, lateral view. USNM 354581. PC, x8. 
9. Same, ventral view. PC, x8. 
10-11. Stereo-pair, internal mold of seed coat, face view, showing band of the raphe traveling along the lower 3/4 of seed periphery. 
USNM 424651. PC, x8. 
12. Same, basal view, with the raphe offset toward one side. PC, x8. 
14 13. Same, rotated 90° from fig. 11. PC, x6. 


LO Stáminate hamamelidaceous inflorescence, UF 5567.05. лы ы en ee e. C NU 50 
14. Medial section through inflorescence in siltstone matrix, showing at least seven florets in oblique-lateral view. Each floret 
set off by perianth and containing conspicuous stamens. Etched, RL, x7. 
15. An individual floret from the same inflorescence sectioned transversely near the apex showing five anthers with interspersed 
small staminodes surrounding clumps of pistillode hairs. Etched, RL, x 14. 
16. Same, sepal in transverse section showing stellate trichomes on the abaxial (left) surface and and simple trichomes on the 
adaxial (right) surface. TL, x65. 


144 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 15 


Figure Page 
5 Stamma hamamelıdaceous inflorescence, conmumusd:- UF S567 e ан nl ee 50 
1. Anther in transverse section from the floret in Pl. 14, fig. 15, showing pollen within the locules. Thin section, RL, x60. 
2. Same, enlargement showing globose tricolpate pollen grains preserved within translucent silica. TL, x 250. 
3. Tricolpate reticulate pollen in situ within an anther. Etched, SEM, x 750 
4. Polar and equatorial views of the pollen, showing reticulate ornamentation and elongate colpi. Etched, SEM, x 1500 
5. Detail of exine showing smooth wall of the aperture and columellae beneath the reticulate tectum. SEM, x 7500. 
6. Рута инв sterile flower irom Gouser Road Clatmo locality 238. UF 9962... се елин кн ie sige a кек anne 51 
ISA И ее A ШИ ЖОШО RS TR Ones И NN те а OE add Куу» RAS нако ae re 50 
7. Fruiting panicle from the Nut Beds showing apparent trifurcate branching and numerous capsules. OMSI Pb697. х2. 
8. Counterpart of same, enlarged to show capsule with two styles, thickened apical disk and longitudinal striations. x8. 
9. Extant Hydrangea arborescens L. Fruiting panicle with persistent fertile flower. Fruits showing longitudinal striations, thickened apical 
disks and two styles each. Summit Lookout Mountain, Tennessee. A. Cuthbert, Sept. 16, 1908 (FLAS 9652). X2. ............. 


15 
PLATE 

58 

UMBER 

ICA AMERICANA, N 

GRAPH 

PALAEONTO: 


ae 


[p 


PLATE 16 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


oa ~ 


Figure 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 145 


EXPLANATION OF PLATE 16 


m EI Ee CMR UW ПОЛ Авр ПО е M p cL о Mrd i i m ре 50 


ie 


Tricarpellate capsule, apical view, showing triangular cast of dehiscence opening. Holotype, UF 6445. SEM, x8. 


2. Tricarpellate capsule, basal view, UCMP 10669. x8. 


12% 
13. 
les: 


. Bicarpellate capsule, basal view, showing wedge-shaped cast of dehiscence opening. UF 6444. x 10. 


Same, basal view, showing thickness of septum. PC, x 10. 


. Same, lateral view, showing apical wedge where the capsule was splitting. PC, x 10. 

. Lateral view of the holotype, cf. fig. 1, showing transverse striations. x8. 

. Basal view of a bicarpellate specimen. Note fingerprint pattern of fine transversely oriented striations. UF 8696. x8. 

. Same, sectioned transversely, showing pair of deeply intruded parietal placentae. Etched, RL, x 10. 

. Same, prior to sectioning, lateral view, showing transverse fibers of the endocarp and remains of the mesocarp composed 


of small polygonal cells and longitudinal fiber bands (arrows). UF 8696. SEM, х 15. 


. Transverse acetate peel section of the tricarpellate specimen in fig. 2, showing deeply intruded, parietal placentae and 


scattered dislodged seeds (arrows). UCMP 10669. TL, x10. 


. Transverse section of a bicarpellate fruit, showing single locule with deeply intruded, parietal placentae and loose seeds. UF 


6446. Etched, TL, x 10. 

Thin section of a fruit showing three seeds. UF 9385. RL and TL, x50. 

Thin section of the same fruit showing seeds with embryo in cross section. RL and TL, x50. 
Isolated seeds from fruit dissolved in hydrofluoric acid. UF 9883. TL, х 50. 


16. Extant Hydrangea anomala D. Don. Seed showing stout hilum, elliptical central body and surrounding wing for comparison with 
figs. 14, 15. Ishikari Province, Japan. K. Miyaper(Mo 8858810 a) RE ай EEE 509 е 51 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 17 


Figure Page 
lj AREY Fa mn ЕА АТОНА oh oe ee ИЙ Оо MES 51 
1. Chalcedony endocarp cast, face view, showing areolate pattern of surface ridges and more or less elliptical outline, UCMP 
10702. РС, X5. 


2. Apical view of same, showing median keel with a raised trough (arrow) representing Ше funicle. РС, х5. 
3, 4. Stereo-pair of endocarp mold in siltstone, face view, showing the characteristic reticulum of surface ridges with funicle 
following fruit contour (arrow). UF 6458. х5. | 
5. Chalcedony locule cast from the same specimen as in figs. 3, 4, face view, showing reticulate surface, median longitudinal 
groove representing path of the raphe, running between the hilum (h) and chalaza (с). PC, х5. 
6. Same, apical view, showing relatively narrow outline and keel in plane of bisymmetry. PC, х5. | 
7. Same, enlarged to show punctate locule cast surface. SEM, х20. 
8. Detail of same, showing rows of circular punctae representing papillae of the locule lining. SEM, х 280. 
с ШКОК ОШ ОШОО ONE cS DL ML DM о IR M ve ort in чуге С е 53 
9. Chalcedony locule cast with very faint surface reticulations. The locule cast is partially broken to show the smooth seed 
cast inside. Note median groove of the raphe (r). Holotype, USNM 424743. PC, x5. 
10. Same, basal view showing smooth contour of the seed, the circular chalaza (с), and groove of the raphe (т). PC, х5. 
11. Same, detail of locule cast surface, showing characteristic punctae. SEM, x 100. 
12. Same, detail of seed cast surface, showing rows of rectangular cells. SEM, x 100. 
13. Locule cast, face view, showing truncate base and asymetrical apex with weak median ridge and very faint reticulum. USNM 
422383. РС, X 5.5, 
14. Same, lateral view, showing pointed apex, truncate base and longitudinal keel in plane of bisymmetry. РС, х 5.5. 
15, 16. Same, basal view, showing the intersection of ridges forming a cross pattern. РС, х 5.5. 


PLATE 17 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 18 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 147 


EXPLANATION OF PLATE 18 


Figure Page 
¡ES ОЛО ООО О AS of salon es us а ols A раб охо ЛЕ c c MN 55 
1. Endocarp, face view, showing longitudinally rugulate and verrucate surface. Holotype, USNM 354788. РС. 
2. Somewhat abraded specimen, face view, showing rough, longitudinally ribbed surface. USNM 354789. 
3. Mold of an endocarp in siltstone matrix, showing veinlike reticulum. USNM 446085. 
4. Endocarp, face view, showing showing reticulate pattern of surface grooves. UF 8618. PC. 
5. Holotype as in fig. 1, apical view, showing roughly lensoidal outline. 
6. Transverse section of specimen in fig. 2, showing elliptical outline of locule, horizontal plane of dehiscence, and funicular canal 
(arrow). 
7-10. LOGICA DORI TIG RISESDUDONS ^B. ун К ta tear tote ae Pe RENE CL O M E. O M eases a 54 
7. Endocarp, face view, showing elliptical outline and relatively smooth surface. Holotype, USNM 354816. 
8. Smaller specimen, face view. USNM 354796. 
9. Apical view of specimen in fig. 7, showing lensoidal outline and course of funicle along the right margin. 
10. Same, transverse section showing thickness of wall and outline of locule, partially filled with chalcedony and quartz crystals. 


"oe 
11. Zodicarpa ampla sp. nov. Transverse section of specimen in figs. 2, 6, showing pappilate locule lining (above) and elongate, sinuous- 
АШЕПКЕ Ое о ea M ch NEM LL n HE у 53 
Ib d Tod cannes ООУ sp env + details опа ша Одо с пати а о = = es 54 


12. Transverse section of holotype, figs. 7, 9, 10, showing papillate locule lining (above) and sinuous-walled cells with intracellular 
spaces of the endocarp. TL, x75. 
13. Transverse section of the specimen in fig. 8, showing papillate locule (above), and full thickness of endocarp with tissue grading 
\ from more or less horizontally aligned cells near the locule to vertically aligned cells distally. TL, х30. 
14. Same, showing thin seed coat, papillae of the locule lining and intracellular spaces within the endocarp tissue. TL, x75. 


148 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 19 


Figure Page 
FI отнета Ерик БЕК СУРЕ novel з 55 
1. Endocarp, apical view showing funicular bulge. Holotype, USNM 354783. PC, х 1.8. 
2. Same, ventral view, showing longitudinal groove leading to Ше funicular bulge. PC, x 1.8. 
3. Same, basal view showing radiating meridional grooves. PC, х 1.8. 
4. Somewhat abraded endocarp, apical view showing the transverse funicular slit; source of sections in figs. 7, 9. USNM 446084. 
XT. 
5. Holotype as in figs. 1-3, lateral view showing longitudinal grooves and the apical funicular bulge. PC, x 1.5. 
6. Dorsal view of an endocarp showing longitudinal groove in plane of bisymmetry, and apical funicular bulge. UF 8596. PC, 
х 1.5. 
7. Same ѕресітеп as іп fig. 4, sectioned transversely, showing thickness ofendocarp wall, and the seed with two thick cotyledons. 
RL, X2. 
8. Transverse thin section of the specimen in fig. 6, showing thickness of endocarp wall. RL, x 10. 
9. Detail from fig. 7, showing balloon-like рарШае lining the locule. RL, x 185. 
ТРУ СИС ИОС SD ONE RN а улуу. a ШАКАЛ мулы т. M T IE 57 
10. Епдосагр, face view, showing ovate outline and punctate exterior. Holotype, UCMP 10708. х 1.5. 
11. Same, basal view. x 1.5. 
12. Same, sectioned transversely, showing thin endocarp wall and elongate endocarp projections into Ше locule. RL, x 1.5. 
13. Same, magnified to show tubular structure of the endocarp projections. RL, x7. 
14. Same, anatomy of endocarp projection showing elongate, undulate-walled cells. RL, x 20. 
LTE съ а авва МЫНЕ (RN ts MEN кумк у ките uii sane лк бе RE O ota диети 51—57 
15. Pyrenacantha zenkeri (sp. Moore) Exell. Transverse section of fruit showing thin endocarp wall with elongate protrusions into 
the locule. Angola. Gossweiler 8730 (BARC). RL, x4. 
16. Phytocrene macrophylla (Bl.) Bl. var. macrophylla (det. Н. Sleumer). Endocarp with typical surface pitting. Leyte Island, | 
Philippines, January 1906. A.D.E. Elmer 7134 (А). x0.9. \ 
17. Phytocrene oblonga Wall. Transverse section showing pits of the endocarp without protrusions into the locule. *Austro Cohui 
Chinae”. L. Pierre 2837 (A). х 2.5. 
18. Iodes cirrhosa Turcz. Papillate lining of the locule. Perak, Ipoh, Malaya. S.C. Chin 994 (L 401157). SEM, x 27.5. 


PLATE 19 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 20 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 149 


EXPLANATION OF PLATE 20 


Figure Page 
o БЫК QOL не ТАСОС Sote ee О S 56 
1. Locule cast, face view view showing prominent pits and obovate outline. USNM 312751. х1. 
2. Same, lateral view, showing thickness and narrowed apex. х1. 
3. Transverse section of permineralized endocarp; some of the wall has broken away from the locule cast. Note thickness of wall, 
hollow protrusions into the locule, and shriveled seed coat. UF 9567. RL, х1.25. 
4. The same, magnified to show detail of endocarp wall including a protrusion into the locule. RL, x 7.5. 
5. Anatomy of the endocarp showing interdigitating cells and tiny papillae lining the locule (top of figure). UF 9567. RL, x60. 
Gala ECLA COD PY LOCKEN CAD SCUG O pers ICSC Ma КЛ en О ОУ a э EC 56 
6. Endocarp, face view, with pitting indistinct at the surface, prior to sectioning (figs. 9, 11). UF 8597. x 1.5. 
7. Endocarp with part of the outer layer broken away revealing numerous pits. UF 8608. х 1.5. 
8. Locule cast with adhering remains of the endocarp. OMSI Pb181. x 1.5. 
9. Same specimen as in fig. 6, lateral view, showing thin profile (contrast with fig. 2), and position of transverse section shown in 
fe [Чу 169: 
10. Same, basal view, showing cross-shaped intersection of longitudinal ribs. x 1.5. 
| 11. Same, transverse section showing relatively thick wall, hollow protrusions of endocarp into locule, and collapsed seed coat. 
| 9.9. 
| 12. Locule cast, face view, showing prominent pits and wide, nearly circular, outline. UF 8599. х1.5. 
13. Same, lateral view, showing thickness and apical Кее]. х 1.5. 


| 
| 
| 


150 


Figure 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 21 


E ИБ ШО УО о ИИ ERR МО л к ИИ ЗООС MOEN И TON 


1. 


2; 
3. 


8. 
9. 


Chalcedony nut cast, lateral view, showing smooth elliptical outline, small apical projection and meridional grooves. UF 5338. 
РЄ, хз. 

Same specimen, apical view, showing smooth circular outline with about 12 meridional grooves. PC, х 1.5. 

Nut opened along the plane of dehiscence showing thick wall, and the locule with two prominent cavities corresponding to 
the two cotyledons. UF 4872. x2. 


. Permineralized nut, transverse section showing two lobes of the locule, two distinct placental bundles in the primary septum, 


and four flattened lacunae in the nut wall. UF 7965. RL, х2.5. 


. Chalcedony locule cast, lateral view showing extent of primary septum. UF 9461. x2. 
. Same, rotated 90°, showing straight shoulder regions, median line of dehiscence and rounded base of lobes without development 


of a secondary septum. x2. 


. Chalcedony locule cast in lateral view showing position and extent of primary septum, and two cotyledon lobes. UF 9458. 


х2, 
Same, rotated 90°, indicating weak formation of a secondary septum. x2. 
Same, viewed apically, orientation the same as specimen in fig. 4. х2. 


10-17. ОЛИСИ ООШ CLOW I) Manchester, a.d So I Н Felt Л жешс E E «Гм Si Ten e's и e ERE 


10. 


Br 
12; 


13. 
14. 


Transversely compressed samara viewed apically showing four elongate wings radiating from a circular nutlet. OMSI Pb1580. 
Xd 

Smaller specimen with well-preserved nutlet as seen in fig. 14. UF 9612. x2. 

Longitudinally fractured specimen showing elongate pedicel (p) and the wings arising equatorially from the nutlet. OMSI 
Pb899. x4. 

Detail of wing showing subparallel, thin dichotomizing venation. ОМІ Pb776. х 5. 

Detail of nutlet from the samara in fig. 11, sectioned transversely showing plane of dehiscence, oriented vertically, and three 
lobes of the locule; a fourth locule lobe, upper right, is not preserved but may be inferred from the symmetry. RL, x 10. 


. Transverse section in the basal one-third of an isolated silicified nutlet showing four lobes of the locule, delimited by thick 


primary horizontal and secondary vertical septa. UF 8867. RL, x8. 


. Same, sectioned closer to the equator, near the top of the secondary septum, with the four basal lobes of the locule joining to 


form two lobes. Note lacunae in the wall at both sides of the primary septum. RL, x8. 


. Chalcedony locule cast showing basal cleavage from the primary and secondary septa. USNM 354375. SEM, x10. 


57 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 21 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 22 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 151 


EXPLANATION OF PLATE 22 


Figure Page 
Заир AICO DIAIVCARVO2aHICKEV1 p DOV. ne ee Lp M An EEN 60 
1. Infructescence after dispersal of fruits, lower half showing natural longitudinal fracture, upper half sectioned longitudinally, 
revealing broadly triangular helically arranged bracts. UF 9251. x3. 
2, 3. Stereo-pair, longitudinally fractured permineralized infructescence, showing stout axis and intact locule casts (arrows). Holotype, 
UF 6454. x5. 
4. Same infructescence fractured transversely, showing two bilobed locule casts (arrows) arranged around the central axis (a). 
IS, 
5. Locule cast removed from the same infructescence, showing prominent groove of the primary septum, rounded base of lobes 
and pointed apex. SEM, x 16. 
6. Isolated locule cast, face view, showing cleft from primary septum. UF 8763. SEM, x20. 
7. Same as fig. 5, apical view, showing narrow dorsiventral thickness. SEM, х 15. 
. Same, basal view. SEM, х 16. 
E OF. Palacocary a clarnensis Манса c a ne, 59 
9. Isolated locule cast. UF 8756. SEM, х15. 
10. Permineralized nut, lateral view, with part of the nutshell fractured away, revealing the locule cast with a median cleavage 
corresponding to the septum. USNM 424640. PC, x 10. 
11. Same, rotated 90°, showing acute base and median longitudinal rib. PC, x 10. 
12. Same, apical view, showing rounded outline and single longitudinal rib (lower part of figure). PC, x 10. 
13. Same specimen, sectioned transversely near the equator showing thin fruit wall, incomplete primary septum and shriveled 
seed within the bilobed locule. RL, x 15. 
14. Nut, lateral view, showing elongate-conical base of nut. Nutshell broken away in upper half, revealing the rounded, apically 
pointed locule cast. Note the prominent median longitudinal rib. UF 9361. SEM, х 10. 
15. Chalcedony locule cast showing smooth contour and median line of dehiscence. Note absence of secondary septum development. 
UF 8752. SEM, x12. 
16. Same, apical view, showing plane of dehiscence (vertical) and septum (horizontal). SEM, x 12. 
17. Nut sectioned transversely near the equator, showing endocarp with line of dehiscence and septum. USNM 458416. RL, x15. 


192 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 23 


Figure Page 
PME SSMO ПИ M den E o E КУ шу. 61 
1, 2. Stereo-pair, ventral view, showing globose shape and faint ridge running from the equator to the apex. USNM 424728. PC, 
NU SD. 
2, 4. Same, stereo-pair, apical view, showing circular outline and ventral ridge. PC, x 7.5. 
5. Same, sectioned transversely, showing thin testa and vertical line separating the large planoconvex cotyledons. RL, x 10. 
6. Specimen, with remains of endocarp adhering to locule cast. Holotype, USNM 435006 PC, x5. 
7. Same, enlarged to show columnar cells of the endocarp. SEM, х 50. 
8. Same, detail of columnar cells. SEM, x 100. 
guys ey OCU PAI CIEE ао DV S cq IM ce m uud LU. rug eU ери DN T 
9. Endocarp, basal view, showing circular outline, smooth surface and prominent attachment scar. USNM 424807. PC, x3. 
10. Same, lateral view, showing height nearly equal to width, truncation at the attachment scar. PC, x3. 
11. Endocarp dislodged from its cupule (fig. 13), basal view showing elliptical attachment scar. Holotype, UF 9406. PC, x3. 
12. Same, lateral view, showing relatively short length. PC, x3. 
13. Same, in matrix with basally attached cupule, longitudinal section. RL, x4. 
14. Same, enlargement of cupule tissue, showing frequent oil cells with contents preserved as dark spots, and showing the contact 


61 


of the cupule with the base of the fruit. RL, x25. | 
15. Same, detail of transition between cupule (c) and seed (s). Note the uniseriate layer of dark columnar cells of the endocarp 
(arrow). RL x50. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 23 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 24 


| 
| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 153 


EXPLANATION OF PLATE 24 
Figure 
ыз, Bauno@urpum NANCOCKIL sp non Ноо ре ОМ УРА а а л л ccc c 62 
1. Lateral view of specimen with adhering siltstone matrix, partially broken showing thickness of wall surrounding locule cast. 
Хо: 
2. Transverse section showing circular outline, unilocular with shriveled remains of the seed. х2. 
3. Fruit wall showing columnar layer lining the locule and outer parenchymatous layer with stone cells. х 30. 
den Bauroeanpummuibedensis sp. nov. Molotype USNM 435090. 200000000000 62 
4. Lateral view showing more or less elliptical outline and smooth surface. The base is incomplete. PC, х5. 
5. Transverse section showing circular outline, thick endocarp wall and remains of two large cotyledons that filled the seed cavity. 
RL, x7. 
6. Same, enlarged to show cells of the endocarp, locule lining and seed coat. RL, х 15. 
7. Same, acetate peel, showing endocarp composed of parenchyma with occasional interspersed larger, more rounded oil cells, and 
junction with seed coat. TL, x50. 
15. LECGUTOCUND UMN GISINO1G ESAS): ЙОМ ОЮУ ОБОО И а ал c a a 62 
8. Basal view, showing central attachment scar, some adhering siltstone on the lower left. х 5. 
9. Lateral view, showing fine longitudinal grooves. x5. 
10. Same, showing transverse plane of section used for fig. 11. x5. 
11. Transverse section, showing thick wall and unilocular, single-seeded construction. RL, x8. 
12. Detail from transverse section, showing numerous oil cells with dark contents scattered through the mesocarp. RL, x 50. 
13. Detail of wall layers adjacent to the locule (lower part of figure). Note locule lining of uniseriate thin-walled columnar cells, 
next a 5—8-seriate layer of sclereids, followed by a tissue of large parenchyma cells with occasional interspersed oil cells. RL, 
x 200. 
14. Same, showing exocarp of columnar cells exterior to the mesocarp tissue of parenchyma cells with interspersed oil cells. RL, 
x 200. 
15. Mesocarp tissue showing several closely spaced oil cells. RL, x 270. 


8- 


154 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 25 


Figure Page 
(SAA о SURE I oe COSE нар АЕ ЗА Cc Lr и оси И 
1. Fruit with numerous seeds in siltstone. UF 5675. х8. 
2. Same, transverse section showing seeds in various orientations. RL, x 10. 
. Isolated permineralized fruit, transversely broken. UF 5668. x 15. 
Same, transverse section, showing angular to rounded seeds with white embryo casts. RL, x20. 
. Longitudinal section showing persistent pedicel, hypogynous calyx (с), longitudinal septation (s) and radiating seeds. UF 
6449. RL, x10. 
6. Detail from fig. 4, showing several seeds intercepted at levels showing the enlarged columnar cells of the germination valve 
(v). RL, x40. 
7. Transverse section of a seed from the specimen in figs. 1, 2, showing the embryo cavity (e), and the ventrally thick layer 
of isodiametric cells through which the raphe (r) passes. RL, x 50. 
8. Transverse section of another seed in the same fruit. UF 5675. RL, x50. 
9 SE Don TOCONDOT sp. OMSIT PHI T23. AL Suecus ure no nee лкнн кыыл Esser а erae su оа es dr diss e e DL EDS 63 
10-12. Magnolia tiffneyi sp. nov., chalcedony seed cast. Holotype, UF 6549. PC, хб. 255 у/........1...... rr enn 65 
10. Ventral view, showing median raphe groove and deep cleft at heteropyle with protruding stalk. 


11, 12. Dorsal view showing V-shaped moat with stalk extending longitudinally. | 


PLATE 25 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PLATE 26 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 155 


EXPLANATION OF PLATE 26 


у Figure Page 
ОНОЛ Агар A E | ee а ЕИ I И 64 
1. Infructescence permineralized in sediment, transversely fractured showing thick woody folicles; note seed upper left. 
Holotype, UF 6529. х3. 
2. Same, transverse section with four folicles radiating from the central axis. x3. 
3-9. Chalcedony seed casts. PC, x 6. 
3. Ventral view, showing ovate outline, pointed hilar end and median basal raphe groove. USNM 424698. 
4. Dorsal view, showing concave heteropyle area. UF 6556. 
5. Smaller specimen, ventral view, showing well preserved heteropyle. UF 6546. 
6. Broader specimen, dorsal view, showing slight collar at the chalaza end. USNM 424697. 
7, 8. Stereo-pair of same specimen, ventral view, showing slight swelling or collar surrounding a small moat and stalk. 
Viewed in stereo, the stalk is seen to extend ventrally in contrast to the straight stalk of other species (cf. Plate 25, 
figs. 11, 12). 
10-17 9. Another specimen, dorsal view, showing faint collar at the base. UF 6555. 
ell. Magnolia paroblonea sp. nov., chalcedony seed casts. PG, ХА o ee ee 65 
10. Dorsal view, relatively wide specimen. UF 9188. PC, x6. 
11. Same, basal view showing elongate chalazal groove. PC, x6. 
12, 13. Stereo-pair, narrower specimen showing pointed hilar end and truncate base. Holotype, UF 9192. BOX 
14, 15. Same, stereo-pair, basal view showing elongate chalazal groove. PC, x6. 
16. Ventral view, with elongate chalaza groove at base and longitudinal striations. UF 9191. PC, x4. 
17. Same, basal view showing wide heteropyle groove. PC, x4. 
18. Magnolia muldoonae sp. nov. Basal view of the specimen in fig. 5. PC, хб. но нненненненненне нн nn 64 
Ka Magnolia tiffneyi sp. nov. Basal view of the specimen in Pl. 25, figs. 10-12. PC, X6. нн rr rr 65 


Magnolia muldoonae sp. nov. Striations on the surface of chalcedony seed cast from fig. 9. SEM, х40...................... 64 


156 


Figure 


1-7. 


8,9% 


10-20. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 27 


Page 
AT INLET МОО а ЗОО e le re O Are Та ер ИР 66 

1. Endocarp showing smooth surface, dorsal keel and basal pedicel scar. USNM 424641. PC, x5. 

2. Cross section of specimen in fig. 1, approximately equatorial, showing thick endocarp wall, central bilobed condyle sur- 
rounded by locule with remnants of seed coat. х5. 

3,4. Successive serial longitudinal sections separated by 2 mm. х5. 
3. Longitudinal section intercepting lateral periphery of the bilobed condyle showing circular endocarp wall and locule 
outline. UF 8728. 
4. Longitudinal section closer to the medial plane, showing bilobed condyle, horseshoe shaped locule and remnants of 
seed coat (arrow). 

5. Longitudinally fractured endocarp, showing cordate condyle with median septum and surrounding subhemispherical locule. 
Holotype, UF 8745. х5. 

6. Median longitudinal section of an endocarp in matrix, basal portion, showing sediment-filled apertures of the condyle. UF 
6308. x7. 

7. Longitudinal section showing anatomy of pericarp, with fiber orientation grading from periclinal on the inner part of the 
endocarp to anticlinal on the outer 2/3 of the endocarp. Some parenchyma cells of the mesocarp are preserved, top of 
photo. UF 5712. TL, x100. ‘ 

Extant Anamirta cocculus (L-J. Wight & Am. JIN. Gowanloch (BARC). Xd За 5... cec i ee ce ieee bene ee nn 66 

8. Fruit sectioned through the condyle, cf. fig. 3. 

9. Fruit sectioned in medial plane, showing bilobed condyle, horseshoe-shaped locule and bumpy endocarp surface, cf figs. 

4, 5. 
СОО ЖАШОО У Е а Au cub IU И SN, кудаи, aor Агалина Ulin A С А 66 
10. Endocarp with faint median line of dehiscence. USNM 354472. x1. 
11. Laterally fractured specimen showing locule surrounded by thick-walled endocarp with rounded lacunae. UF 9249. x1. 
12-14. Cast of the lacunae system in the endocarp. UF 6850. х1. 
12. Ventral view, showing outline of the condyle, and four projections representing the apical and basal apertures. 
13. Dorsal view showing median cleavage corresponding to suture line of the endocarp wall. 
14. Lateral view showing reticulation of lacunae system. 
15. Apical view showing medial dehiscence line and paired apical apertures opening to the system of lacunae. UF 9254. x 1.29. 
16. Transversely fractured specimen, showing crescent-shaped locule, paired ventral condylar cavities апа lacunae within the 
endocarp. USNM 354537. х 1.5. 
17-20. Босше casts. РС, X 1:25. 
17. Ventral view showing longitudinal median keel and subapical placenta (arrow). UF 6844. 
18. Same, dorsal view, showing median keel, pointed apex and base. 
19. Ventral view of a larger locule cast showing pointed apex and base. UF 6845. 
20. Same, dorsal view showing faintly irregular surface. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 27 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 28 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 157 


EXPLANATION OF PLATE 28 


Figure Page 
С Ооа Aue a een T E 67 
1-5. Chalcedony endocarp cast. USNM 424850. PC, x 2.5. 
1. Dorsal view, with median longitudinal keel and scabrate surface. 
2. Ventral view showing condylar cavity with faint transverse striations and median keel. 
3. Apical view showing the pair of cavities. 
4. Basal view showing bilobed cavity. 
5. Specimen with portion of endocarp broken away revealing thick wall and locule cast. UF 6869. x3. 
6-9. Chalcedony locule casts. PC, x 2.5. 
6. Dorsal view, showing median keel and flattened base. UF 6862. 
7. Ventral view of same, showing median keel, transverse striations and subapical placental scar (p). 
8. Lateral view. UF 6863. 
9. Same, ventral view, showing shorter length relative to fig. 7. 
10. Endocarp in transverse section showing thick wall surrounding the crescent-shaped locule. UF 6873. RL, x4. 
CUY COCOM ChASSICTUSTOC: SDN ON В CL er ИОА totes ОЕ 69 
11. Endocarp cast, dorsal view. USNM 424849. PC, x 2.5. 
12. Same, ventral view of condyle cavity with median keel. PC, х 2.5. 
13. Dorsal view of locule cast showing median keel. OMSI Pb471. PC, x3. 
14. The same, ventral, showing cup-shaped cavity and median keel. PC, х3. 
15. Apical portion of locule cast with some adhering endocarp. Holotype, UF 6769. PC, x3. 
16. The same, cross section, showing the thick endocarp surrounding the locule. RL, х3. 
17. Apical view of the specimen in figs. 13 and 14. PC, x3. 
18. Extant Calycocarpum lyonii (Pursh) Gray. Transverse section showing relatively thin endocarp wall. Locule similar to that in 
НЕМ С MTOR dauGUris5934 (A): З Е 70 


11-17. 


158 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 29 


Figure Page 
1. Calycocarpum crassicrustae sp. nov. Transverse section of holotype illustrated in Pl. 28, figs. 15, 16, enlarged to show cells of the 
endoeatpewallanditesta TL бО A eter eer uitae eor Mu US ES У ЕНЕ НЕ 
2-8. OS PONE CIONES MOVE Use. IE URS у deine e eMe tar cU E Ip SUM IUE Enid DU Manet ER SUD ATL HE 
2. Locule cast viewed dorsally, showing median keel and smooth surface. Holotype, USNM 354502. PC, x4.8. 
3, 4. Stereo pair of the same, viewed ventrally, showing condylar cavity, median keel, and a thin layer of adhering endocarp at 
the lower right. PC, x4.8. 

. A larger locule cast, ventral. USNM 424648. PC, x5. 

. Apical half of a broken locule cast. USNM 424649. PC, x4.8. 

. Stereo-pair of the same specimen, looking at the transversely broken end, showing the rounded condyle cast with a median 
groove representing the median keel of the ventral wall of the endocarp, surrounded by the c-shaped locule cast. The thin 
film between the locule cast and the condyle cast indicates that the endocarp was thin-walled. PC, x 4.8. 

OT SUO DO NINA Na РАТОВАО NOV RR ә оста Туз» NE CO EN OR Real ВАН ие ыйы NIE DOT CUT A TIT МЕМ ки та ыл 
9, 10. Stereo-pair of chalcedony locule cast, dorsal view, showing median keel and apical point, lower half of locule cast broken 
and removed, showing mold of ventral wall of the endocarp in sedimentary matrix. Holotype, UF 9255. x6 
11. Mold in siltstone matrix of the ventral surface of an endocarp, showing impression of the medial keel, supapical placenta 
and spines near the base. UF 9256. SEM, x6.5. 
12. Same, showing protrusion of endocarp spines into the matrix (arrows). SEM, x10. 
13. Chalcedony locule cast, ventral view, USNM 326715B. x6. 
1426 E КО КУО Sp: NOV... cux Ep cM o UE eds quedo aud does а пати НА te pa EL MEET f ts 
14. Sedimentary locule cast, lateral view, showing smooth circular outline, central plug and cleavage where the apex and base 
meet. USNM 446075. PC, x4. 
15. Locule cast viewed ventrally, showing median keel, and cleavage where the apex and base of Ше endocarp meet. Holotype, 
USNM 354506. PC, x4. 
16. Same, viewed laterally, showing central plug. x4. 


70 


о ON tA 


7, 


70 


74 


| 
| 
| 
| 


PLATE 29 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PLATE 30 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Figure 


1-5, 


6-10. 


11-14, 


15-20, 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 159 


EXPLANATION OF PLATE 30 


HNO о ове ти sp- nov: Holotype USNM 422383. pee "xu ccu c Fi 
1. Locule cast in lateral view, showing thin profile. 
2, 3. Stereo-pair, dorsal view showing pointed apex and base and median keel. 
4, 5. Stereo-pair, ventral view with shallow condylar cavity, prominent median keel, and subapical placental scar (arrow). 
Лаан рО OON DEW ESPOSA er. 71 
6. Locule cast in lateral view showing thin profile. 
7, 8. Stereo-pair, dorsal view showing shallow pits and very faint median keel. 
9, 10. Stereo-pair, ventral view with pronounced pitting and prominent median keel. 
Dimeus arani ОЙУ (НОО) СОЕ ООУ К see 73 
11. Edge view of a locule cast limb protruding from matrix, showing characteristic sculpture, surrounded by the dark outline 
of the endocarp mold. Note the three-angled apical rim of the endocarp. UF 6485. 
12, 13. Stereo-pair of horseshoe-shaped locule cast and carbonaceous endocarp remains in matrix, showing fluted rim and central 
foramen. UF 6484. 
14. Locule cast with one limb broken, showing placental “knob” and evenly spaced fluting. UF 6486. 
ECIGCOSINOMICNIUMUVEnUD est Chandir PE oa. а a ee еы ыл T 74 
15. Mold, face view, in siltstone showing endocarp ornamentation. UF 9545. 
16. Edge view of an endocarp cast. USNM 354561. 
17, 18. Stereo-pair of same showing numerous flutings along the rim. USNM 354561. 
19. Locule cast showing smoother contours than do the endocarps (figs. 17, 18) and pronounced sinus between the uneven 
limbs. USNM 354562. 
20. Larger locule cast with more asymmetrical limbs. UF 6487. 


160 


Figure 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 31 
Page 


155. Curvitinospora formanii gen. et sp. nov. Holotype, USNM 326714. РС, -X 8. er. Hu. d eee ann a Доу + 71 


ЈЕ 
2.034 
4, 5. 


Dorsal view of locule cast, showing prominent median keel, rounded base and pointed арех. 
Stereo-pair, ventral view, showing longitudinal depressions on either side of the median raphe ridge. 
Stereo-pair, lateral view with ventral surface to the right, showing longitudinal curviture. 


6-9. Thanikaimonia geniculata gen. et sp. nov. Holotype, USNM 355480 PC, Х6.5. 2... rane ec анан ee тен кина 75 


6. 
1, 8. 
p. 


Face view of lateral limb, showing smooth surface, basally tapered profile, and longitudinal median keel. 
Stereo-pair, lateral view, showing curvaiure. 
Basal view, showing attachment scar and unequal development of the limbs. 


10-14. Dayisicarpum limaciordes sp: nov. Holotype, USNM 435054 2... eh nu ee ere lente aptos tee egret з 73 


10,11. 


Stereo-pair, permineralized endocarp, lateral view showing curved dorsal ridge with fine radially oriented striations and 
concave base of the endocarp partially embedded in sediment. RL, х7. 


. Same, edge view with plane of bisymmetry medial, showing median dorsal keel and the convex lateral margins of the fruit, 


best seen on the left side. RL, х7. 


. Same, sectioned transversely at right angles to the plane of symmetry, showing diamond-shaped locule at top of specimen, 


endocarp walls that bulge and thicken around the condyle. RL, x 10. 


. Same, showing detail of seed coat within locule and thin surrounding endcarp wall. RL, x40. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 31 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 32 


Figure 
1-14, 


17-19. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 161 


EXPLANATION OF PLATE 32 


Dinseieroresonense:Manchesienelsktessi u ie ee К, 

1. Apical view of permineralized seed showing round profile and cicular hilum. Holotype, UF 6621. PC, х5. 

2. Lateral view of seed with fibrous seed coat partially broken away exposing smooth endocarp cast, upper right. UF 6627. PC, 
x5. 

3. Lateral view of seed with portion of seed coat broken away and exposing the silica infilling of the central chamber. UF 6611. 
Re: 

4. Internal mold of seed coat showing the prominent micropylar plug with protruding beak. UF 6623. PC, x5. 

5. Apical view of the same specimen, showing intact plug. PC, x5. 

6. Lateral view of the specimen in fig. 1 showing striate seed coat and apical truncation from which Ше operculum has detached. 
РС х5, 

7. Basal view of the specimen in fig. 2 with chalazal mass exposed through the broken seed coat. PC, x5. 

8. Basal view of the specimen in fig. 3, showing chalazal mass and ribbing of the seed coat. PC, x5. 

9. Basal view of the specimen in fig. 4, showing position of chalaza. PC, x5. 

0. Specimen with micropylar plug removed. OMSI Pb1314. PC, x5. 

1. Median longitudinal section of the holotype in figs. 1, 6, showing silica-filled central endosperm chamber separated by a basal 
septum from the chalazal mass and by and apical septum with a central pore from the operculum. RL, x5. 

12. Median longitudinal section of the specimen in figs. 2, 7, showing two-layered seed coat, lower right, and intact embryo at the 
apex of the central chamber. RL, x5. 

13. Detail of the embryo from fig. 12. x8. 

14. Detail of the specimen in fig. 11, showing two-layered seed coat. х8. 


. Extant Ensete glaucum (Roxb.) Cheesman. Seed sectioned longitudinally for comparison with figs. 11—14. Marobe District, Papua 


INE Wa оү T E ук а ei c ны та Shine, SIM MN MILL. NM E 76 
Ensete oregonense seed cast in siltstone showimgantact apical operculum, OMSI.PbI0G6I RS. 005.0 oo. ne T5 
Detail of the specimen in fig. 10, showing construction of the seed coat. n у л a л 73 


17. Oblique apical view of the inner mold of seed coat showing concave surface with central circular hilar scar. SEM, х 15. 

18. Lateral view showing longitudinally striate seed coat partially broken away from the endosperm cast, revealing smooth inner 
surface. SEM, x50. 

19. The same, apical view, showing relative thickness and stratification of seed coat. SEM, x75. 


162 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 33 


Figure Page 
IG Saba bDIachNeNens&(Chatidien);Marzseedeasis 2's ee ee 76 
1. Lateral view of seed showing raphe groove and protrusion of embryo. UF 8487. PC, х6. 
2, 3. Same, stereo-pair, ventral face showing circular depression with central raised chalazal mound and raphe groove. PC, х6. 


AS Sava СТРИПОВЕ Та Ре РЈ Соор ао seed.easis..... a Ри IOTER Н E qe АР и SG ОНИ 77 
4. Ventral view, showing Ше сисшаг depression, central chalazal mound, and raphe groove. UF 9751. РС, х5. 
5. Lateral view showing groove and chalazal mound. OMSI Pb1812. PC, х5. 
6. Same, ventral view showing chalazal mound within circular depression. PC, х5. 
7. Specimen showing elliptical pad of tissue surrounding the chalazal mound. PC, x5. UF 9752. 
8. Transverse section showing bi-layered seed coat. UF 9754. RL, x65. 7 


ОДИ отр СОВЕ Manchesten N RE ER. А Е Е ДИЕ 
9. Infructescence broken transversely showing radiating florets of mature fruits and the receptacular impressions indicating five 
free carpels per floret. Holotype, UF 5153. RL, x3. 
10. Tangential section of silicified infructescence, showing five-fruited florets. UF 5159. Etched, RL, x5. 
11. Transverse section of an infructescence, showing florets, with fruits and perianth in longitudinal section. UF 5158. RL, х4.5. 
12. Same specimen as Figure 10, detail of a fruitlet in transverse section showing layers of pericarp and surrounding the locule 
with a single seed. TL, х 50. 
13. Platananthus synandrus Manchester, staminate inflorescences possibly from the same plant as M. glabra. Inflorescence sectioned 
medially showing florets with stamens enveloped by well-developed perianth. UF 5151. x7. ..............................: 78 
14. Platanus hirticarpa sp. nov., medial section of infructescence, showing numerous radiating florets. Holotype, UF 5160. x7. ... 78 


PALAEONTÖGRAPHICA AMERICANA, NUMBER 58 PLATE 33 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 34 


Figure 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 163 


EXPLANATION OF PLATE 34 


Page 


1-4. Platanus CANDO sp- nov- Continue inom JL По Holotype ШЕ ЗЛЕ ee 78 


ЈА 
2: 
3. 
4. 
Tanyoplatanus cranei SUN СПЕЦ О на TERN на LL AME и RA tud iE EE Gn Red. А die EUR E 78 
5. Longitudinally broken infructescence in matrix, showing achenes attached to central axis. UF 5671. x4. 

6. 
- Same, median longitudinal section through the axis with rows of achenes on either side. Note protruding perianth segments. 


5-15. 


Oblique peripheral section, showing perianth, hairs and individual achenes. RL, x20. 

Longitudianal section through two achenes with obtriangular outlines with the surrounding perianth of a single floret. RL, x 20. 
Longitudinal section of an achene with typical persistent style. RL, x 20. 

Longitudianl section through the dispersal hairs of an achene. RL, x 50. 


Same, transverse section showing circular central axis with radiating achenes. RL, x8. 


RE x8. 


- Tangential section of the infructescence showing three- and four-carpeled florets. RL, х8. 

- Median longitudinal section showing clusters of at least two carpels. RL, х8. 

- Detail from fig. 8, showing three closely adhering carpels surrounded by perianth segments. RL, x20. 

- Same, with four closely adhering achenes. RL, x20. 

- Longitudinally fractured infructescence larger than that in fig. 5. Holotype, USNM 424876. x3. 

- Same, tangential section of infructescence showing three- and four-carpeled florets. х 16. 

- Same, transverse section of fruits, enlarged to show dispersal hairs distributed between the achenes. x 50. 

- Same, oblique-longitudinal section of achenes including one with elliptical seed preserved in white silica. x 20. 


164 


Figure 
1-10. 


11-15. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 35 


OS на аи АН STONER Е NO какас a LN РАТ К MEE let БИ 
1. Locule cast, lateral view, showing pyriform shape. UF 6461. РС, х5. 
2, 3. Stereo-pair, base, showing thin layer partially broken away revealing circular chalaza. UF 6460. PC, х5.7. 
4. Same, lateral view. PC, x5.8. 
5. Same, apical view, showing apical keel. PC, х 5.6. 
6, 7. Stereo-pair, specimen with adhering testa and sedimentary matrix, showing circular chalaza and raphe groove. Holotype, UF 
9262 РС, X5. 
8. Same specimen, transverse section showing elliptical outline and thickness of the endocarp wall. PC, x6. 
9. Same, thin section showing locule lining surrounded by sclereids. RL, x75. 
10. Same, detail of endocarp wall showing sclereids with scattered rhomboidal crystals (arrows). TL, x 300. 
РИП еи. аро ШЕШУ pe ШЕ ООШОТ vn + othe CORSI DD E 
11. Lateral view of permineralized endocarp showing keel in plane of symmetry, ovate outline and gently verrucate surface. PC, 
х4. 
12. Face view showing ovate-elliptical outline, and thicker funicular margin, right side. РС, х4. 
13. Transverse section showing thick wall with plane of dehiscence near bottom of figure in plane of bisymmetry. RL x6. 
14. Same, enlarged, showing partially detached locule lining, left side of figure. RL х25. 
15. Detail of sclereids composing the endocarp. TL, х 100. 


80 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 35 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 36 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 165 


EXPLANATION OF PLATE 36 


Figure 


=н, EIN ЛОО UA р ом Uc D Rome a E ord Mu T ERE 80 


1. Infructescence with elongate, pedicellate capsules. Scar at arrow indicates the position ofa detached fruit, suggesting opposite 
arrangement of fruits. OMSI Pb1307b. x1. 
2. Infructescence with portions of three attached permineralized capsules. The one on the right is nearly complete. Note absence 
of perianth scars at the junction of the pedicels with the fruits, indicating inferior ovaries. Holotype, OMSI Pb1197b. x2. 
3. Detail of capsule from the counterpart of specimen in fig. 2, showing longitudinal median septum. Holotype, OMSI Pb1197a. 
X215; 
4. Transverse section of a capsule from the specimen in fig. 2, showing two carpels with axile placentation, and relatively thin 
pericarp. RL, x8. 
5. Transverse thin section of the same, showing a T-shaped placenta, surrounded by numerous radially arranged seeds. TL, 
127 
6-11. Acetate peels of permineralized fruit. UF 9333. 
6. Transverse section showing well preserved seeds surrounding collapsed placenta, and a portion of remaining pericarp. 
TE 12. 
7, 8. Detail of individual seeds in transverse section, each with a central body surrounded by а thin dark layer, which is in 
turn loosely enveloped by the seed coat formed ofa reticulum of cells with projecting anticlinal walls, and thin, mostly 
collapsed, outer periclinal walls. TL, x 50. 
9. Longitudinal section through a seed. TL, x30. 
10. Longitudinal, peridermal section of a seed, showing reticulate seed coat. х25. 
Е 11. Detail of seed coat showing reticulate thickening of cells composing the seed coat. x65. 
> 13. Extant Emmenopterys henryi Oliver. Southwest China. С. Forrest 26041 (US 1377648). нон 81 
12. Isolated seed showing dark elliptical central body surrounded by thin reticulate patterned wing. x 10. 
13. Detail of seed coat showing reticulate thickening of cells composing the seed coat, cf. figs. 10, 11. x 25. 


166 


Figure 


1-5. 


10-12. 


13-15. 


16-19. 


ОООО рО TC SUE SE MOSER EIN EN ee ER О ee e 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 37 


Page 
луда асите ареала ar ANS AAA ee AR Who ee о, 82 
1. Endocarp, ventral view, showing angular outline, circular funicular plug, rounded basal flank. UF 6702. 
2. Same, lateral view, showing curvature of dorsal surface. 
3. Locule cast, ventral view, showing funicular opening and median longitudinal keel. Note punctate surface. Holotype, UF 
6673. 
4. Same, basal view, showing punctate surface and median keel. 
5. Same, lateral view showing height (vertical dimension) similar to length (horizontal dimension). 2 
6. Endocarp, ventral view, showing prominent funicular cavity, rounded-triangular outline and relatively flat basal flank. UF 
6715: 
7. Endocarp, ventral view, with funicular plug and relatively flat basal flank. Holotype, UF 6714. 
8. Sarne, dorsal view, showing median longitudinal keel and rounded outline. 
9. Same, lateral view, showing rounded dorsal surface and truncate ventral side. 
Menos clonadas ono ИО ОНУ Dese NIETOS ee ЛЫ ыу иж у н UN ИЕ 
10. Ventral view showing rounded sides, median keel and circular funicular plug. 
11. Same, basal view (cf. fig. 4), showing funicle, median keel and height much greater than width. 
12. Same, lateral view (cf. fig. 2). 
MOTOS aj CH ei elie Reidel: сапер ы put qu REN ER IE ee ne 
13. Ventral view of a locule cast with some endocarp adhering (upper right), showing large overall size and large funicular 
plug. UF 6706. 
14. Same, dorsal view, showing smooth locule surface and median longitudinal keel. 
15. Locule cast, ventral view, with some of the endocarp preserved in the funicular area. UF 6707. 
Ме ИЕНЕН ОРАК да ре drove Holotype OE 072 РО 0r c Tas кук wie ii aM o her Ae eC ву A DR 
16. Ventral view showing circular funicular plug, median keel and narrow width. 
17. apical view showing height much greater than width. 
18-19. Stereo-pair, lateral view. 


82 


84 


83 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 37 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 38 


11-16. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 167 


EXPLANATION OF PLATE 38 


Page 
- Meliosma endocarps in longitudinal section in the plane of bisymmetry. RL, х 5.5............... а. 82 
PMG OST GM CUSEKOIMUASD ENO UU T ee. 82 
EOS ТРОН ВР ато OB OVID rr р MEER et 82 
ESOS MORE ЕТА ПА СТАЕ СКА ШЕШ БОВ o су ко. 84 
< Melosma leptocarpa эр. nov., longitudinal section. UF 6739. un 83 
MOON verda (Decker) comi aon, РС, SS. 0 5 0 O зз ы кууз LEM a 84 
5. Endocarp cast, lateral view, showing cratered surface and ventral notch. UF 6729. 
6-7. Stero-pair, lateral view. Note funicle scar within the ventral notch. UF 5245. 
8. Same, viewed from base, showing circular funicle scar, endocarp thickness and median keel in the plane of symmetry. 
EE EOOD К ТОРОО ТОУ SEME НА en 86 


9. Chalcedony embryo cast, lateral view, showing globose outline, convoluted cotyledons and curved radicle. UF 6589. 

10. Face view of cast, showing wide elliptical, nearly circular, outline, and the U-shaped tip of the radicle. Holotype, UF 6587. 
Ша СО ODN ОСОО SD ROME ууу. уу-уу . (ADM улык c. TOOT а hee 86 

11. Chalcedony embryo cast showing convoluted cotyledons and elongate radicle. USNM 424668. SEM, x8. 

12. Embryo cast with portion of seed coat adhering, upper right. UF 6596. SEM, x8. 
13, 14. Stereo-pair, chalcedony seed cast with portion of testa flaked away showing convolutions of embryo cast and basal protrusion 

of the hilum. Holotype, USNM 424669. PC, x8. 

15, 16. Stereo-pair of same, basal view. Note hilar projection near the center. PC, x8. 


168 


Figure 
1-6. 


7-16. 


17-21. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 39 
Page 


DIEBE ONC Л ТОЛГА MOV a ae ао ee 85 


ЈЕ 


чл bh шо ~ 


Samara showing broad lateral wing with arching dichotomous venation and elliptical endocarp with straight margin of 
abscission aligned with the stipe. Holotype, OMSI Pb1133. x 2.2. 


. Similar fruit. OMSI Pb567. x2.2. 

. Larger specimen, showing style (s) arising from the elliptical endocarp. OMSI Pb638. x2.2. 
. Specimen showing reticulate venation over the endocarp. OMSI Pb1786. x2.2. 

. Specimen with narrower wing. OMSI Pb802. x2.5. 

6. 


Well preserved smaller specimen with recurved style from Clarno White Cliffs locality. UF 9859. x4. 


Витара ОООО Уо: Internal molds of: ie seed coat а ere peres MM кз varese s dert су. ж АИ 86 
7. Lateral view, showing smooth, rounded base and sides and the oblique apical truncation. USNM 424663. PC, x6.5. 


15, 16. 


СИЛИ ЗИ РАНОИІС Вр AON SECO GAS ee e e кердк na e RD ee уы ы beber ee ED TT A E о. 

. Lateral view of seed cast showing smooth surface, rounded base and oblique apical truncation. USNM 424681. PC, х 6. 
. Same, apical view, showing rounded-triangular outline, rounded rim of the hilar truncation, and central hilum. 

. Holotype, lateral view. UF 8743. PC, x6. 

. Same, ventral view. 

. Same, apical view, showing obovate truncation with central hilum from which veins radiate, with a fragment broken away 
near the micropyle. 


. Same, stereo-pair, apical view, showing circular hilar truncation, and position of micropyle (m). PC, x6.5. 

. Translucent seed cast, ventral view, showing median longitudinal course of the raphe. USNM 424662. RL, x6.5. 

. Same, apical view, showing emergence of raphe at center of hilar truncation. RL, x6.5. 

. Lateral view showing smooth rounded sides and oblique truncation. Holotype, USNM 424644. PC, x6. 

. Same, stereo-pair, apical view, showing raphe that passes from ventral (lower) surface to the apical truncation and adhering 


vascular reticulum. PC, x6. 
Same, ventral view, showing median longitudinal course of raphe. PC, x6.5. 


PLATE 39 


ka 
kd» 
4: 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 40 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 169 


EXPLANATION OF PLATE 40 


Figure Page 
ТЫ адла ENS A eR TOU О оо л 87 
1. Complete seed cast showing veruccate surface and hilar sinus with bulging lip. UF 6577. x8. 
2. Seed cast (internal mold of tegmen), ventral view. USNM 424654. x6.5. 
3. Same, lateral view, showing indented hilar region. PC, x 6.5. 
4. Seed with tapering micropylar limb and deep hilar region. Holotype, USNM 424655. PC, x 7.5. 
5. Same, tilted to show path of raphe along dorsal surface. SEM, x 10. 
6. Same, ventral view. SEM, х8. 
7. Same, sectioned dorsiventrally, showing layer of columnar cells making up the seed coat, and dorsal position of the raphe 
(arrow). RL, х 15. 
8. Seed cast (internal mold of tegmen). UF 6581. RL, x6.5. 
9. Same specimen as 2, 3, sectioned longitudinally. RL, x 10. 
10. Same specimen as in fig. 8, sectioned longitudinally. RL, x8. 
iaig 11. Same, showing uniseriate columnar layer of the tegmen. RL, x 20. 
РЕВА И menge se ЭДЕ АЫ. MIT nor ДЕ, TS 88 
12. Schisandra chinensis Baill. Seed with smooth surface. Shinano Province, Japan, Sept., 1960. Miyoshi Furuse (A). x8. 
13. Schisandra viridis A.C. Smith. Seed with rought surface and hilar sinus with bulging lip. Cf. fig. 1. Chekiang, China. Ren- 
Chang Ching 5132 (A). x8. 
14. Same, sectioned longitudinally, x 11. 
15. Kadsura japonica Dunal. Longitudinally sectioned seed. Ohobu, Kobe, Japan. Н. Muroi 6569 (А). RL, x30. 
16. Schisandra chinensis. Same as fig. 12, sectioned longitudinally. RL, x10. 
|. Kadsura japonica, sectioned longitudinally, same as fig. 15. RL, х10. 


- Schisandra viridis, sectioned longitudinally. Note columnar inner layer of seedcoat, and proliferation of isodiametric cells of 
the outer seed coat filling the hilar sinus. Same as figs. 13, 14. x 20. 


170 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 41 


Figure Page 
Ea occidentalis Manchester. ЕНА ЛАРИ 88 
1. Chalcedony fruit cast, ventral view, with features of the seed expressed through the thin wrinkled pericarp, including the 
central circular chalaza, raphe, and pedicel scar (top of figure). UF 5201. PC, х7.5. 
2. Same, dorsal view, showing rugulate ornamentation. RL, X 7.5. , 
3. Ventral view of fruit cast with verrucate surface layer partially broken away revealing internal mold of seed coat with 
prominent circular chalaza at the center and pointed micropyle at the apex. Holotype, UF 5200. PC, х7.5. 
4. Permineralized fruit in matrix, ventral view. UF 9283. RL, x7.5. 
5. Internal mold of seed with successive layers of seed coat and pericarp adhering. UF 9692. SEM, х 10. 
6. Same, cellular detail on surface of internal seed coat mold. SEM, x 150. 
7. Median lateral section of the specimen from fig. 4, showing thickness of fruit wall and seed coat layers, c-shaped endosperm 
cavity conforming around the chalazal complex (right side). UF 9283. RL, x 10. 
8. Transverse section, showing endosperm within the seed. UF 9284. RL, x10. 
9. Same, transverse section enlarged to show anticlinally oriented cells of seed coat. RL, x 50. 
ОА AS AA A m dE Se uoo Haec qiue o ot c MM ва dE 89 
10. Chalcedony endocarp cast, lateral view, showing truncate apex, rounded base and meridional grooves. Holotype, USNM 
354596. РС, x3. 
11. Same, basal view, showing central attachment scar surrounded by radiating meridional grooves. PC, x3. 
12, 13. Stereo-pair of same, apical view, showing four apertures on the apical truncation. PC, x3. 
14. Another specimen, showing the apical truncation and meridional ribs, broken below. USNM 354603. PC, х3.8. 
15. Same, apical view, showing at least two apertures within the truncation. PC, x 3.9. 
16. Chalcedony endocarp cast, lateral view. USNM 354592. PC, х5.7. 
17. Same, basal view. PC, х5.9. 


PLATE 41 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


2 29 


Зенодот 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 42 


6-15. 


16-25, 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 174 


EXPLANATION OF PLATE 42 


Page 
CARA A re ee E T A NU E UU E 89 
1. Seed, lateral view, showing reticulate seed coat. Holotype, ОЕ 6501. SEM, x15. 
2. Same, basal view showing thickness of seed. SEM, x16. 
3. Internal cast of seed coat, lateral view, showing embryo curving around the condyle. UF 9449. SEM, x15. 
4. Internal cast of seed coat, showing condyle in center and possibly the micropylar limb on the left, UF 9450. SEM, x19. 
5. Detail from holotype, fig. 1, showing thick anticlinal walls of cells forming the seed coat. SEM, x 50. 
EAS AS О ку S MS ДА REN ERN rl cS нос I E ors лш. 90 
6. Silicified endocarp, apical view, showing rounded contour, median ridge and elliptical obliquely oriented funicular pore. The 
median ridge is deflected laterally at the funicular opening. Holotype, USNM 355693. PC, x5. 
7. Same, lateral view, showing wide-elliptical outline and verrucate surface. PC, x 5. 
8. Endocarp, apical view, with obliquely oriented funicular pore. Lower portion of endocarp wall broken away exposing smooth, 
rounded locule cast. USNM 424629. PC, x5. 
9. Chacedony locule cast, showing smooth surface and well-marked longitudinal keels that intersect at the funicular pore. UF 
VB PO х5. 
10. Same, lateral view showing relatively narrow elliptical outline. РС, х5. 
11. Locule cast, apical view, surrounded by remains of Ше endocarp wall. USNM 422384. SEM, x 7.5. 
12. Some of the endocarp wall removed from the specimen in fig. 11, showing layer lining the locule. SEM, x10. 
13. Longitudinal section of a well preserved endocarp fragment showing dark wall of endocarp surrounding coiled embryo. UF 
9328. RL, x8. 
14. Section of wall from the holotype, figs. 6, 7, showing outer layer of prismatic cells including crystaliferous inflated cells. RL, 
x 20. 
15. Detail of wall from fig. 13 showing large columnar cells. TL, x 60. 
(GE ITISSDUTMIATHICSSD:nONE Lus Reo х=. бош иссе DR ONU ME M CA MR I 9] 
16. Endocarp, apical view, showing nearly circular outline, reticulate surface pattern and four longitudinal ribs. Holotype, UF 
9600. PG, X6. 
17. Same, lateral view, showing median longitudinal rib, reticulate smaller ribs. PC, x6. 
18. Same, rotated 90°, showing primary longitudinal rib and acute apex. PC, x6. 
19. Same, basal view showing intersection of four median ribs. PC, x6. 
20. Locule cast showing smooth surface, circular chalaza, and radicle at top of figure. USNM 355429. PC, x6. 
2. Same, lateral view, showing curved radicle. PC, x6. 
22. Longitudinal section of permineralized specimen, showing single locule filled by the embryo which has a curved, protruding 
radicle. Note thickness of endocarp and subapical position of funicular plug. UF 9581. x10. 
23. Transverse section of fruit, showing large, nearly circular locule surrounded by successive layers of the pericarp. UF 9581. 
x 10. 
24. Detail of same, with locule to the left showing succession of pericarp layers, the most prominent of which is the uniseriate 
layer of square crystaliferous cells. RL, x 15. 
25. Detail of internal surface with digitate cells. Seen by focusing down through the outer translucent part of the pericarp. UF 
8626. RL and TL, x35. 


192 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 43 


Figure Page 
ЗЕЕ cp EN SO uL D MEL M M M UH IUE EU 92 
1. Silicified endocarp, face view, showing median longitudinal keel, apical attachment scar and veruccate surface ornamen- 
tation. UF 8658. SEM, x6. 
2. Same, lateral view showing apical notch, rounded outline and typical surface ornamentation. PC, x6. 
3. Lateral view of an endocarp with part of the wall fractured away, revealing locule cast with prominent curved radicle. UF 
8641. SEM, x6. 
4. Transverse section of an endocarp. Horizontal line is the separation between radicle and the rest of the embryo. UF 8651. 
RL, x10; 
5. Same, enlarged to show wall anatomy. RL, x30. 
O е аСТ рО ood NET NU m. co ao TN иеа Lr O AS DEC ME 92 
6. Silicified endocarp, dorsal view, showing median longitudinal keel, ovate outline, and relatively smooth surface. Holotype, 
UF 9781. SEM, x15. 
7. Same, lateral view. RL, x 10. 
8. Same, sectioned longitudinally, showing coiled embryo of the seed, with pronounced radicle. RL, x 10. 
9. Transverse section, showing thickness of wall and horizontal line separating the radicle from the rest of the embryo. UF 
9776. RL, x10. 
10. Same, enlarged, showing columnar cells of endocarp wall. RL, x50. 
11-13. Isolated locule casts. 
11. Dorsal view showing median keel. UF 6792. PC, x10. 
12. Ventral view. UF 6793. PC, x10. 
13. Lateral view. UF 6792. PC, x10. 
14. Longitudinal section, showing complexity of the apical region. UF 9585. x10. 
15. Same, showing columnar cells of the endocarp wall. x 18. 92 


16-182 Gon пеи те ти esquete ns Мао не Кер <ar i ri e rr sud ave merces taedia euam чуу. uestri gan c ае 
16. Typical samara showing large wing with veins converging toward the stigmatic area, and small wing forming a crest along 
part of the endocarp. OMSI Pb565. x4.5. 
17. Smaller samara. OMSI Pb1502. x5. 
18. Fruit showing the small wing extending parallel to the long axis of the endocarp. UF 9285. x6. 


PALAEONTÖGRAPHICA AMERICANA, NUMBER 58 PLATE 43 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 44 


Figure 
1—5. 


6-10. 


11-15, 


16-18, 


2 
22 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 173 


EXPLANATION OF PLATE 44 


И пи по UN ee se ee Lo tI к 96 
l. Chalcedony seed cast, ventral view, showing short parallel infolds, and prominent median groove along the raphe ridge. 
Holotype, UF 6534. PC, x 7.8. 
2. Same, dorsal view, showing spatulate chalaza and chalaza-apex groove. PC, х8. 
3. Dorsal view of a seed cast with much of the seed coat flaked away revealing internal mold of the seed cavity with circular 
chalazal mark. UF 6533. PC, х8. 
4. Chalcedony internal mold of seed coat, ventral view, showing sharply pointed base and parallel infolds. UF 6535. PC, x8. 
5. Same, dorsal view showing chalaza-apex groove. PC, x8. 
ООН ТООС veu ED ER САН m ИИ © s 94 
6. Chalcedony seed cast, ventral view, showing short divergent infolds, and median groove along the raphe ridge. UF 6537. PC, 
x8. 
7. Same, dorsal view, with portion of seed coat flaked away, revealing smooth internal mold of the seed cavity. Note chalaza- 
apex ridge (contrast with chalaza-apex groove, figs. 2, 3, 5). PC, x8. 
8. A broader seed cast with widely divergent infolds. Holotype, UF 6536. PC, x8. 
9. Same, dorsal view, showing spatulate chalaza, chalaza-apex ridge and chalaza-base groove. PC, x8. 
10. Chalcedony internal mold of seed coat, ventral view, showing pointed base, sharp raphe ridge, and prominent, divergent 
infolds. UF 6538. PC, x8. 
IDE СЕ ООО аро ма e A ari. s to Mua E cq up Mq ER ш. 94 
11. Seed cast, ventral view, showing cordate outline and short, divergent ventral infolds. Holotype, USNM 424665. PC, x8. 
12. Same, dorsal view. PC, x8. 
13. Internal cast of seed cavity, ventral view, showing prominent short ventral infolds and sharply pointed base. UF 6542. PC, 
x8. 
14. Same, lateral view, showing split along dorsiventral plane. PC, x8. 
15. Same, dorsal view, showing elliptical chalaza and shallow chalaza-apex groove. PC, x8. 
AURA LISS CTT А SD, MON S CE D UTER соо у е, os cc RM E үт E 93 
16. Chalcedony internal mold of seed coat, ventral view, showing width and depth of infolds and cordate outline. UF 6543. PC, 
хб. 
17. Same, dorsal view, showing elliptical chalaza and chalaza-apex and basal groove. PC, x 6. 
18. Chalcedony internal mold of seed coat cavity with corresponding external mold of the same seed showing ventral infolds as 
а pair of mounds in the siltstone matrix. Holotype, UF 6572. x 3.75. 


- Vitis tiffneyi sp. nov. Transverse section showing deep ventral infolds and relatively narrow seed coat. UF 9573. RL, х12.5.. 96 


20. Same, enlarged greater to show columnar cells of seed coat. RL, x 30. 
Ampelopsis rooseae sp. nov. Transverse section showing relatively thick seed coat. UF 9575. RL, х12.5.................... 94 
Ampelocissus auriforma sp. nov. Transverse section showing broad ventral infolds and thick seed coat. UF 6571. RL, x6. ... 93 


174 


Figure 


1-5. 


14-17. 


NIE UAEM A E AOS AEE И 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 45 


АО СЕН Вр HO л ED e е сн ОГЫ 
1. Chalcedony seed cast, ventral view, showing long divergent infolds, relatively flat ventral faces meeting at the raphe ridge. 
Some of Ше seed coat is broken away apically, exposing internal cast of seed coat. Holotype, UF 6539. PC, x8. 
2. Same, dorsal view, showing smooth rounded surface and oval-spatulate chalaza. PC, x8. 
3. Another specimen with long divergent infolds. UF 6540. PC, x8. 
4. Same, dorsal view, showing spatulate chalaza. Lacking a groove between chalaza and base. PC, x8. 
5. Transverse section. UF 9583. RL, x 12.5. 


" Prmenocaistansusasiicar Scott. UUSINIVEAS 4986" PE On 55 


6. Ventral view, internal mold of seed coat, showing cordate outine with сјећ at apex and long divergent infolds. 
7. Dorsal view, showing circular chalaza and prominent chalaza-base and chalaza-apex grooves. ob 
8. Chalcedony internal mold of seed coat, ventral view, showing rounded apex and long divergent infolds. USNM 355098. PC, 
хб. 
9. Same, dorsal view, showing circular chalaza, radiating shallow grooves and more pronounced chalaza-apex groove. РС, x6. 
10. Largest specimen, showing long ventral infolds. USNM 355127. PC, x5. 
11. Same, dorsal view showing circular chalaza and well-developed chalaza-apex groove. PC, x 5. 
12. Another specimen, showing flattened ventral face and fine radiations. USNM 434985. PC, x5. 
13. Same, dorsal view, showing obcordate chalaza with pronounced chalaza-apex groove and shallow radiating grooves. PC, x5. 
ANDI MOCO GO vo dect olla Clasp Noy Holotype, USNM 353614. enese s OUS oet eR DOE T RES 
14. Lateral view, showing elliptical outline, with a portion of the pericarp broken away revealing striated inner layer. RL, x 1.5. 
15. Transverse section, showing single locule surrounded by thin endocarp and exocarp. RL, x 1.5. 
16. Same, showing detail of seed tissue, and endocarp, including a thin dark layer representing the locule lining. RL, x 12.5. 
17. Same, more greatly magnified, showing anticlinally oriented columnar cells of the locule lining, surrounded by a thick layer 
of poorly preserved schlerenchyma. RL, x50. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 45 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 46 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 175 


EXPLANATION OF PLATE 46 
Figure 


Page 
Les, PIU ОРОЛ ЛОО ее ер апо EU NN CCS ах Ора ie c M ие E ы ea 97 
1. Oblique ventral-lateral view, showing rounded base, pore near the apex, and apical crest. Holotype, UF 6290. SEM, x20. 
2. Ventral view, showing narrow width, median apical crest and pore. USNM 424712. SEM, x20. 
3. Lateral view of a specimen that has broken apically, showing protruding inner cast. UF 9799. SEM, x20. 
4. Same, magnified to show thickness of wall and smooth surface of inner cast. SEM, x42. 
5. Seed sectioned transversely, showing elliptical outline of inner chamber and longitudinal slit leading to the ventral pore. UF 
9800. TL, x30. 
6-10. ESTOS NAAA AMAS NONE ee a ee 59 
6. Fruit showing руг огт shape and scabrate surface. UF 6320. SEM, x 10. 
7. Fruit with part of the pericarp fractured away revealing globose locule cast. Holotype, UF 6318. SEM, х 10. 
8. Locule cast showing globose shape, circular chalazal scar at base, and slight apical swelling. UF 6343. SEM, x 20. 
9. Locule cast, apical view, showing circular outline. UF 9770. SEM, x20. 
НЕ 10. Longitudinal section of fruit in fig. 6, showing globose seed inside, х 10. 
E IA LEE SS DECALS (PLN. OL БЛАТА E A tnt EU И Eu LLL i pM СЕТЕ 97 
11. Specimen viewed laterally, showing J-outline. UF 9808. x15. 
12. Lateral view showing J-outline, neck-like constriction at top. Holotype, UF 9806. SEM, x20. 
а 13. Same, face view. UF 9807. SEM, х20. 


апета оао оеп себер пот SEM K 16 rt. a ne, 99 
14. Ventral view, showing median longitudinal slit. Holotype, UF 9766. 
15. Same, dorsal view, showing truncate base and pointed apex. 
16. Ventral view, specimen with fractured apex. UF 9767. 
y Same, dorsal view showing veruccate surface. 


176 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 47 


Figure 
I IDONEOS УРАН СОК РОМЕ sp. DOVT ee a E o a ee ы E 
1-4. Chalcedony locule cast. Holotype, USNM 424642. PC, хб. 
1. Face view, showing rounded scar at the base, gentle median ridge and protruding apex. 
2, 3. Stereo-pair, same specimen reverse side. 
4. Basal view, showing thin profile and prominent circular scar. 
5-7. Chalcedony locule cast with adhering remains of pericarp. UF 9404. 
5. Dorsal view, showing pointed apex and rounded base. SEM, x6. 
6. Same, tilted back slightly and enlarged to show circular chalaza at base. SEM, х 15. 
7. Detail of adhering pericarp. Punctate appearence is due to parenchyma-like cells, the outermost layer having flaked 
away. SEM, x40. 
8. Transverse section of a permineralized fruit with seed showing two horizontally oriented, parallel cotyledons. UF 9471. 
RE, x15. 
9. Transverse section ofa locule cast with shriveled embryo inside consisting of two parallel, horizontally oriented cotyledons. 
USNM 424643. RL, x15. 
10. Transverse section of pericarp, detail from fig. 8, showing outer layer of columnar cells, inner layer of isodiametric cells. 
RL, x30. 
PESTO ESOS K AA ИСО A S Ле AONE gate ы ыйы ERU emn e Зх 
11. Ventral view, showing rounded sides, truncate apex and two ventral faces joining in a median keel. UF 9366. PC, x8. 
12, 13. Same, Stereo-pair, lateral view, showing convex dorsal side and relatively straight ventral side, disruption of tissue at the 
apical truncation. PC, x8. 
14. Ventral view, specimen tilted forward to show triangular truncation at the apex. Holotype, UF 6392. SEM, x8. 
15. Same, apical view, SEM, x8. 
16. Apical view of an asymmetrical specimen. UF 9364. SEM, х 10. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 47 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 48 


en 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER БАТ 


EXPLANATION OF PLATE 48 


Figure Page 
E Comminicarpa friisae СПЕВ ТО AA E EH AE XEM Le оа а IMMER М о А 98 
1. Longitudinally fractured infructescence. Holotype, USNM 355697. x2.5. 
2. Same, sectioned longitudinally, tangential to the axis, showing closely adjacent fruits in transverse section. RL, x 5. 
3. Same, higher magnification, showing fruits grouped in threes. RL, x 10. 
4. Another infructescence sectioned longitudinally, showing fruits in transverse section that are subangular through crowding. UF 
NSF RE ок Пн 
5. Same infructescence as figs. 1-3, transverse section showing fruits in longitudinal section. RL, X 6.5. 
6. Same, showing detail of individual fruits including persistent style at apex and single seed per fruit. х 8.5. 
7. Detail of fruits from fig. 4, showing the seed coat of a single seed within the locule and pericarp containing a middle layer of 
ig large rectangular cells. x55. 
- Durocarpus cordatus Bene CHSD то У LOLOL DCm UE ШО qm MEER merge Ee c redu s ио быу уло. 99 
8. Permineralized endocarp fruit, dorsal view, showing cordiform outline. PC, x6. 
9. Apical view, showing smooth, rounded contours. Horizontal line resulted from cutting longitudinally for fig. 10. PC, x6. 
10. Longitudinal section through the two locules, showing vascularization of the base and central axis, and thick fruit wall. RL, 
x 10. 
11. Same, enlarged to show tissue surrounding the locule. Acetate peel, TL, x30. 
1% Same, enlarged to show the small, thick-walled endocarp cells lining the locules and the larger, thin-walled cells making up the 
bulk of mesocarp tissue. Note the placental tissue arising from Ше apical end of the ventral side of the locule (arrow). TL, x50. 
13. Transverse section of the same specimen, reassembled after sectioning longitudinally, showing the cross section of locules and 


thickness of septum. Acetate peel, TL, x 10. 


Figure 
1-15. Ferrignocarpus bivalvis gen. et sp. nov 
. Impression of an endocarp valve in sediment, showing thickness of wall and shape of locule cast. UF 9390. x6. 


ps 


2-4. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 49 


Locule cast showing typical smooth surface, USNM 355348. x5. 
2. Face view showing elliptical outline, median longitudinal groove and slightly emarginate base. 
3. Lateral view showing keel in plane of symmetry with apex being slightly more acute than base. 
4. Apical view showing lensoidal outline. 


. Locule cast with adhering remains of endocarp, apical view. Holotype, USNM 424624. PC, x6. 
. Permineralized endocarp in matrix, broken longitudinally showing thick endocarp wall and locule with remains of seed. UF 


ӨЗӘ жэ, 


. Isolated locule cast, face view, showing emarginate base and somewhat broken at apex. USNM 424803. x5. 

. Stereo pair of same specimen in fig. 5, face view showing cordiform locule cast with adhering endocarp. PC, x6. 
. Transverse section of specimen in fig. 2. TL, х5. 

. Successive transverse sections of specimen in fig. 7, showing remains of shrunken seed in locule. RL, х5. 

. Transverse section of specimen in figs. 5, 8 and 9. 


13. Section near the equator showing partial remains of endocarp, shrunken tissue within the locule and the septum. Х 8.5. 

14. Section closer to Ше apex showing incomplete septum. x 8.5. 

15. Transverse section of the fruit wall, with locule at lower side of photo, showing two different sclerenchymatous layers. 
x 60. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 49 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 50 


Figure 
1-4. 


5-13, 


14-19, 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 179 


EXPLANATION OF PLATE 50 


Page 
I PONE NEL ан БЕНО МАНИ NE она a ада rn D LU M TINI qu e 100 
l. Lateral view, showing D-shape outline, and coarse, fibrous ribs. UF 9557. PC, x10. 
2. Same rotated 90*, showing hilar end and lensoidal outline. PC, x 10. 
3. Another specimen, lateral view, showing rounded contour and stright hilar margin to the left. Note the network of bifurcating 
surface ribs. Holotype, USNM 424729. SEM, x20. 
4. Same, rotated 35°, showing the hilar area around which the surface ribs are interrupted. SEM, x 20. 
О Са DIT еугеати та pen. CESP NONE Sec ede OO NO MED миз аы AA Ace LP NU UP cx e 101 
5. Lateral view, showing smooth surface, obovate outline with pedicel projection at base. Holotype, USNM 355663. PC, x7. 
6. Same, basal view showing circular outline and central pedicel scar. PC, x 7. 
7, 8. Stereo-pair of another specimen, lateral view, showing pedicel projection and rounded contours. USNM 435009. PC, x7. 
9. Same, basal view of pedicel scar. PC, x 7. 
10. Larger specimen, lateral view, showing smooth surface and rounded outline. USNM 43501. PC, х7. 
11. Transverse section of the specimen in figs. 5, 6. RL, x 10. 
12. Specimen in fig. 10, sectioned longitudinally, showing thickness of middle layer and globose, smooth walled locule. RL, x8. 
13. Same as fig. 11, close-up, showing the middle layer composed of isodiametric parenchyma cells and the thin inner layer 
surrounding the locule. RL, x 50. 
B TD MIT A eb ode UNION bois cede н abad CD е 100 
14. Seed, face view, showing elliptical seed body with encircling wing. Wing partially fractured away on left side. UF 6286. SEM, 
х 13. 
15. Seed, lateral view, showing longitudinal wing. UF 9867. SEM, x15. 
16. Seed in face view with partially intact wing. USNM 435058. x 10. 
17. Seed in face view, showing the surrounding wing which has broken in places along the left side, and showing the aril-like 
structure near the apex. Holotype, USNM 435057. x 10. 
18. Same, median transverse section, showing wing at top of figure, broken remains of the wing at bottom of figure, and elliptical 
outline of endosperm within the inner seed coat. RL, x 20. 
19. Same, showing two distinct layers of seed coat, and cellular makeup of the endosperm. RL, х 50. 


180 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 51 


Figure 
Fa ел театра ies Halli: eeu врео УУ Ноор УМА А ОВ М СЕ ИИО 101 
1. Lateral view, showing prolate outline and longitudinal grooves. РС, х8. 
2. Transverse section near equator, showing six radially arranged, more or less elliptical locules surrounding a prominent 
central axis, with each locule having one seed. RL, x 10. 
3. Transverse section near the apex, near the ends of the locules. RL, x 10. 
4. Detail of seeds within the locules, transverse section. RL, x 20. 
58. ОООО E И oosa genT IP NOVE О PERS ООВ ИО И PIRE Rees 
5. Basal view of endocarp, showing two valves, venose surface and nearly circular outline. х5. 
6, 7. Stereo-pair, lateral view, broken to show thick wall, thin layer lining the smooth locule and longitudinal keel within the 
locule. х5. 
8. Transverse section showing large polygonal cells making up the endocarp wall, and poorly preserved cells lining the locule. 


x 30. 
102 


Пето OI Se CES под а ee еа а re Ne 
9. Stereo-pair showing pyriform shape and coarse, longitudinally striate surface. Holotype, UF 6111. х5. 
12. Stereo-pair of same, other side, broken and revealing pyriform locule. 
13, 14. Stereo-pair of another specimen showing longitudinal ribs and striations, UF 6578. PC, x5. 
15. Transverse section of same specimen showing thick endocarp wall and single locule. RL, x 10. 
16. Close-up of the same, showing fibers composing Ше endocarp wall, with interspersed clusters of smaller vascular elements. 
Rig, x25: 


PLATE 51 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PLATE 52 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 181 


EXPLANATION OF PLATE 52 


Figure Page 
1-12. ТАНАТ СШТЕП CESDINOV een a 103 
1. Endocarp, lateral view, showing rounded margins and arched striations parallel to the outline ofthe dorsal margin. Holotype, 
USNM 435020. x 7.5. 
. Same, dorsal view showing seed thickness, bisymmetry. х 7.5. 
3, 4. Stereo-pair lateral view, showing arched striations. USNM 422527. PC, 7.5. 
5. Endocarp with relatively smooth surface and faint striations. UF 6340. PC, x7.5. 
6. Endocarp, ventral view, showing circular attachment scar. USNM 435027. х7.5. 
7. Specimen with endocarp partially fractured away revealing curved locule cast. USNM 435045. x 7.5. 
8. Locule cast. Lateral view, showing dorsal curvature and faint ruminations. USNM 435029. PC, x5. 
9. Same, end view, showing longitudinal keel. PC, x5. 
10. Longitudinal section of the specimen in fig. 5, showing crescent shaped seed cavity. RL, x 10. 
11. Longitudinal section, showing large condyle, and seed coat. USNM 435034. TL, x10. 
13. 12. Detail from fig. 11, showing cells of the wall adjacent to condyle, and the seed coat. ТІ, х 25. 


16. Omsicarpium striatum gen. et sp. nov. Holotype, ОМІ РЬ265...... 242и шшк ee nene 104 
13. Lateral view showing ovate outline and surface striations. x 1.5. 
14. Same, rotated 90 degrees. х 1.5. 
15. Basal view. х 1.5. 
16. Tranverse section showing single locule and thick wall. x2. 


182 


Figure 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 53 
Page 


Тез Nm ОД ОУ Юе рей ОН AD e e E лета ON OPES 102 


IK 


Dhu 55) 


Lateral view showing more or less circular outline, persistent pedicel and fine, wavy surface striations. Holotype, UF 8483. 
xI.5. 


. Same specimen, basal view, showing circular outline and radiating striations. x 1.5. 

. Same, transverse section, showing circular locule and thick wall. RL, х 1.5. 

. Smaller specimen, broken to reveal circular locule cast. USNM 354386. x 1.5. 

. Equatorial transverse section showing construction of the fruit wall. Arched line at base of figure is the locule lining. Holotype; 


UF 8483. RL, x10. 


. Same, Transverse section through the mesocarp region. x15. 
. Same, near the periphery, showing transition from mesocarp to exocarp. x15. 
. Same, Detail of the mesocarp, showing anticlinally oriented elongate cells. x65. 


III CODA A MS E DIC узы ту ned cue exa сы E mou EE UE MT UTR DE. ME 103 


. Ventral surface of seed showing dentral depression, radiating striations and protruding radicle. USNM 355414. PC, x8. 

. Same, dorsal view, showing raised circular area chalaza (?) and coarse striations. PC, x8. 

. Stereo-pair of same, viewed laterally, showing raised chalazal (?) area slightly curved longitudinal axis. PC, x8. 

. Chalcedony embryo cast, showing characteristic “screw eye" morphology. UF 5242. PC, x8. 

. Ventral view, with smooth rounded base and shallow central depression, and protruding cast of the radicle. Holotype, UF 


5244. P6, х8. 


. Same, dorsal view. PC, x8. 
. Longitudinal thin section ofa seed showing relatively thick seed coat composed mainly of large rectangular cells, surrounding 


a looped seed cavity with an island of tissue composed of smaller compact cells. Section is slightly oblique and therefore 
does not show the complete loop of the embryo. UF 6346. TL, x15. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 53 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PLATE 


54 


Figure 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 183 


EXPLANATION OF PLATE 54 


Page 
5 Pirtachoides tmata genet sp; Nov. Holotype USNM 122887 PO XI a een 105 
1. Dorsal view, showing ovate shape, median longitudinal keel and faint diagonally arranged striations. 
2. Ventral view, showing continuation of the keel, rounded base and pointed apex. 
3. Lateral view rotated 90° from fig. 2, showing faint surface striations oriented diagonally from upper right to lower left. 
4. Basal view, showing rounded rhomboid outline, central attachment scar and the two valves meeting to form the keel in the 
plane of bisymmetry. 
5. Top view. 
ОЗУИС ОЛЛО ве ешр DON БО ee О EAM 104 
6. Lateral view, showing reticulate sculpture, rounded apical and lateral margins and hilar notch. USNM 355052. RL, х15. 
7. Lateral view showing rounded outline, prominent notch with hilar scar (h) and micropylar point (m). Holotype, UF 6500. 
SEM, x15. 
8. Same, ventral view, showing typical narrow thickness, details of hilum and micropyle. SEM, x15. 
9. Lateral view of a specimen with portion of outer layer broken away revealing underlying cells. USNM 424746. SEM, x20. 
10. Surface detail from the specimen in fig. 6. SEM, x30. 
11. Lateral view of a seed cast with portions of surface reticulum preserved. USNM 424685. PC, x15. 
12, 13. Same, stereo-pair, ventral view. RL, x15. 


184 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 55 


Figure Page 
E PA ООЛО О ОШИ ЖИКА SD TON e еа EN 104 
1. Lateral view, showing ovate outline and typical reticulate sculpture. UF 9605. SEM, x 44. 
2. Apical view, showing circular outline and meridional rows of cells. Holotype, UF 9604. SEM, х42. 
3. Same, oblique-lateral view, showing narrower band of cells (right) that may represent course of Ше raphe. SEM, x40. 
4. Another specimen, abraded apically. SEM, x 40. 
EN РШЕ ШЕ ООО рита рей Ө роу leu lye. EA ES eM a чәр келш ыс ж МИ Ted s ит ва es ENE 104 
5. Internal mold of seed coat, viewed ventrally showing wide-obtuse apex, rounded base and circular “сар”. Holotype, UF 6606. 
SEM, x20. 
6. Another similar seed, viewed ventrally. UF 6425. SEM, х20. 
7. Detail from fig. 5, showing a faint coarse rectagular reticulum superimposed on smaller polygonal cells. SEM, х 52. 
8. Detail of the basal end of the specimen in fig. 5, showing the circular cap. SEM, х 50. 
ES M SCIVIT C сок ee ge DIM NN 105 
9. Internal mold of seed coat, lateral view, showing uniformly angled apex, rounded base and basal cap. Holotype, USNM 424751. 
SEM, x 20. 
10. Another specimen, of shorter length. USNM 424755. SEM, x20. 
11. Same, basal end, enlarged to show cells of seed coat and circular cap. SEM, x62. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 55 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 56 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 185 


EXPLANATION OF PLATE 56 


Figure Page 
Тез Rolygranaenurbedensa.gen. Met ПО О ese осн Rom. К mL Жолу кх ла. Ee. xu. T 106 
. Permineralized fruit, lateral view prior to sectioning, with apex broken. Note widely spaced longitudinal ribs. UF 5665. х4. 
2. Permineralized fruit, lateral view prior to sectioning, with part of pedicel and perianth disk persisting at base, showing elliptical 
outline and smooth surface. Holotype, USNM 355493. x4. 
3, 4. Same, stereo-pair, basal view, showing circular perianth disk and faint longitudinal ribs. x4. 
5. Same, transverse section, showing thin pericarp, and single locule with numerous ovoid seeds in parenchymatous tissue. RL, 
x8. 
6. Transverse section of the specimen in fig. 1, showing single large locule with numerous seeds. Etched, RL, x8. 
7. Fracture surface through a permineralized fruit, showing rounded surface of seeds embedded in parenchyma. UF 5673. SEM, 
х 15. 
8. Section of Ше same specimen, showing the peripheral regions of two neighboring seeds. Note the adjacent parenchyma cells 
and the finely interwoven cells of the seed coat. RL, x 100. 
OI ROMO TOS DEN ОРСОК 105 
9. Lateral view showing reticulate surface and rounded margins meeting to form an angle at hilum. USNM 355448. SEM, х 50. 
10. Same, face view showing apical dorsal keel. SEM, х 50. 
11. Larger specimen, lateral view. Holotype, UF 9830. x50. 
12. Same, enlarged. SEM, x 100. 
[SS бє Pruniticarpa cevallosii peh et Pano AO lO y pe UF 9209, nn 106 
13. Endocarp, lateral view, showing longitudinal rib of carpel closure. Transverse line is the plane of section seen in fig. 15. x 1.5. 
14. Same, base, showing circular outline, pedicel scar and a single longitudinal rib. PC, x 1.5. 
15. Same, transverse section, showing thick wall of endocarp with suture at top of the figure, and single locule with remains of a 
thin seed coat inside. RL, x2. 
16. Same, enlarged to show cells composing the endocarp and the suture horizontally oriented. RL, acetate peel, x 20. 


E 


186 


Figure 


1-3. 


13-19. 


20. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 57 


КООШ MODO ОНИ Oeil, CLUS [ye OVS, Э. ee ee. un КОСЕ: 106 
1. Specimen showing wing with straight upper margin and convex lower margin, and elliptical endocarp. OMSI Pb1381. 
2. Specimen showing three or more sepals arising from the endocarp, right side, and wing with three strong subparallel veins. 
Holotype, OMSI Pb1018. 
3. Well-preserved specimen from the Clarno White Cliffs locality, showing acutely pointed distal wing margin, and two style 
arms protruding from the endocarp. UF 11696. 


b Prnmimperma manumi pen. et Sp. NOV., Seca cdots SEM, ar ea. + «+ wr e е: 107 


4. Lateral view, showing rounded base, pointed micropylar projection. Holotype, UF 9740. 
5. Lateral view. UF 9741. 

6. Lateral view. UF 9742. 

7. Face view, showing narrow width, apical keel and hilum. UF 9743. 

8. Lateral view. UF 9849. 


ОСТЕО Ве еер ОУ и M. RU EA E Mitis iue ee носи E анале КҮЗ 107 


9. Chalcedony cast, face view, showing pointed apex and rounded basal truncation. UF 6402. SEM, х 15. 

10. Chalcedony achene cast, lateral view, tilted back slightly to show circular scar at base. Holotype, UF 6463. SEM, x15. 
11, 12. Stereo-pair of the same, achene preserved in translucent silica showing silhouette of the seed cast inside. TL and RL, x 10. 
OUT ACAVG Vel OSA Gre. er. sp. NOV Holotype, ДИО 4805... re aa nen een TOS ER Teen 108 

13. Lateral view, showing rounded subconical shape. х 4.5. 
14, 15. Stereo-pair of base, showing the central pedicel scar and the rounded bulges of the five lacunae. x 4.5. 

16. Transverse section near base, showing five radially arranged slender pyrenes alternating with the lacunae. RL, х5. 

17. Detail of wall, transverse section, showing uniseriate locule lining (right side), rows of vertically oriented fibers and scattered 

circular resin/gum ducts. RL, x24. 

18. Same, note lack of lining bordering lacunae (left side). RL, x 24. 

19. Transverse section showing isodiametric cells of the pericarp. TL, x45. 
Pruniticarpa cevallosii gen. et sp. nov., detail of cells composing endocarp. Locule and thin seed coat are to the right. Holotype, 
(РИВЕР DUO whe ere niger ee que. RR а eee И HAW Ree UR INEO QUIA Куку аи II ЫЙ rotons. КА ана а 106 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 57 


PLATE 58 


NUMBER 58 


P 


PALAEONTOGRAPHICA AMERICANA 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 187 


EXPLANATION OF PLATE 58 


Figure Page 
1298 Scalaritheca biseriata gem ot Sp: Nov. ee A eU o o йазы еы E она ан 109 
1. External view of a fruit with part of the wall fractured away revealing a row of elongate, curved seeds. Holotype, UF 5674. 
x8. 


2. The same, sectioned longitudinally through the row of about even seeds. x8. 
3. Same specimen sectioned longitudinally at right angles to the section in fig. 2, showing median septum, and a row of seven 
seeds. 
4. Specimen fractured longitudinally showing the thick median septum and two rows of about eight seeds. UF 6107. x 10. 
5. An isolated seed from one of the fruits. UF 9603. SEM, x18. 
| 6. Specimen showing the median septum adjoined by a row of intact seeds (right) and the impression of a row of dislodged 
| seeds (left). UF 6281. SEM х10. 
7. Same, light microscopy. PC, х8. 
8, 9. Same, rotated 90°, stereo-pair. x8. 
10-14. Sambuclspermites ruzulosus PEN Ot SPa NOY Mies ще вон es e ne ee a к 108 
10. Group of three seeds in siltstone. OMSI Pb1716. х 12. 
11. Seed cast. UF 6398. х 30. 
12. Seed cast, dorsal view, showing ovate outline and rugulate surface sculpture. Holotype, UF 6424. SEM, x 32. 
13. Same, ventral view, showing low median ridge leading to the hilum (h), and rugulae. SEM, х 32. 
14. Same, lateral view, showing thin profile. SEM, x30. 
15-17. Sphaerosperma riesii gen. et sp. nov. Holotype, UF 9746. .......................... enne 110 
15. Specimen in siltstone matrix, showing globose morphology, wall partially broken away revealing dark silicified inner tissue. 
Transverse line is saw mark indicating the plane of section for figs. 16, 17. x 1.5. 
| 16. Transverse section, showing wall of uniform thickness surrounding disrupted inner tissue. RL, x 1.5. 
17. Section showing detail of wall, locule to the left. Note lining of rectangular to columnar sclereids and uniform tissue making 
up remainder of wall. x40. 


188 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 59 


Figure Page 
лета ае ве И а ePOS aca eel ee 108 
1. Endocarp, lateral view specimen showing obovate outline and warty surface sculpture. Holotype, USNM 355066. х1. 
2. Same, apical view showing circular outline. 
3. Endocarp, lateral view, showing warty surface sculpture, rounded apex apex and nearly flat base. USNM 354201. x1. 
4. Same, apical view, showing lateral compression, elliptical outline. x 1. 
5. Transverse section of holotype specimen in fig. 1, showing circular outline, thick fruit wall and single circular locule. RL, 


x2. 
6. Transverse section, specimen in figs. 3, 4, showing elliptical outline, single locule and stratification within fruit wall. RL, | 
xD. | 
7. Anatomy of endocarp, transverse section, showing files of rectangular cells and oblique fibrovascular traces. UF 8481. RL, 
XIS 
S- S O EEN ON BEN. et-Sp.nov. d Жук eov tH E еи e ee 109 


8. Infructescence in matrix with ellipsoidal fruits radiating from central receptacle. Holotype, USNM 312750. x2.5. 
9. Chalcedony locule cast removed from infructescence in fig. 8, dorsal view showing pointed apex, rounded base and smooth 
surface. X 7.5. 
10, 11. Stereo-pair of Ше same, lateral view. X 7.5. 
12. Same, locule cast, apical view showing convex dorsal surface and concave ventral surface of locule cast. X 7.5. 
13. Transverse section of fruit removed from infructescence in fig. 8, showing crescent shaped locule and ventral infold of the 
condyle. x 7.5. 
14. Transverse section of specimen with intact pericarp, showing crescent shaped locule cast. UF 9523. х 12.5. 
15. Transverse section of specimen with the seed, endocarp and mesocarp tissues intact. USNM 355368. x 10. 
16. Detail of mesocarp from fig. 14. x 65. 
17. Isolated locule cast, ventral view showing central circular placental scar. UF 5238. x 7.5. 


PLATE 59 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 60 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 189 


EXPLANATION OF PLATE 60 


Figure Page 
1268 ISTHANSDERrMALCOLONGDUNGIAAEM. gom. Сыр. DOV с а а Tce Л а ee 111 
1. Seed, lateral view, showing longitudinal striations and rounded outline. Holotype, USNM 424694. PC, х6.4. 
2. Same, basal view, showing circular outline and radiating striations. PC, X 6.5. 
3. Same, apical view. PC, x 6.5. 
4. Apical view of a specimen showing rectangular cells making up the striations and a cycle of about 11 circular scars surrounding 
the apex. UF 8812. SEM, x 10. 
5. Two seeds adjacent in matrix, with a placenta between them. UF 8718. PC, х5.8. 
6. Detail of surface showing one layer of rectangular cells stripped away to reveal another similar surface. USNM 424873. SEM, 
х 15155 ч 
J— 10 Sphenosperma baccatum gen tSp nov. OO pen ЛЕ S006 по ea ee. 110 
7. Transverse thin section, showing nearly circular outline of fruit, and five well developed seeds. RL, х5. 
| 8. Same, enlarged, showing ап undeveloped seed (arrow) between two fully developed seeds. RL, x6.5. 
9. Same, showing T-junction of septum with fruit wall, with a seed on each side of the septum. RL, x20. 
10. Closeup of the seed coat, showing inner layer (right) of isodiametric cells, and outer layer of columnar cells. RL, х 50. 
МЕТА Srockeycaroa dobosa кеп. et sp: шо ТОО De ШЕОЛ a Mc m m M ыт, ры, н, tH 
11. Fruit exposed in matrix, showing spheroidal shape and smooth outer surface. х 5. 
12. Same, sectioned transversely, showing circular outline. RL, х5. 
13. Same, enlarged showing parenchymatous layer partially pulled away from the outer layer. RL, x8. 
14. Detail of the wall showing inner thin layer of anticlinally elongate cells, middle layer of parenchyma, and outer layer of 


anticlinally oriented fibers. RL, x40. 


190 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 61 


Figure Page 
Кей ЧОРО ДЕО AA OV дение N л EM M E eee te 112 
l. Seed, face view, showing rounded base, protruding apex and smooth surface. Holotype, UF 6108. х5. 
2-3. Same, stereo-pair, lateral view, showing thin, wavy profile and characteristic band with a medial groove running from near 
the apex to the base formed by the margins of the two cotyledons. х5. 
4. Detail from same specimen, lateral view showing folded nature of the lateral band terminating near apex. ca. х 10. 
5-6. Stereo-pair, face view. UF 9528. х5. 
7. Same, basal view, showing the marginal band. x5. 
8. Same, face view, showing cell pattern on the surface. x 35. 
9. Same, transverse section, locule to the left, showing anticlinally oriented columnar cells of the inner seed coat. x 50. 
10-1857 ОЛОВО In. oregontensisuben, О ЫК ION ee ат ырл жне ыу We ы н A ты 112 
10. Chalcedony endocarp cast, lateral view, showing pointed apex, elongate outline, and longitudinal grooves between carpels, 
with some of Ше wall flaked away exposing smooth locule cast. Holotype, USNM 424671. PC, х8.5. 
11. Chalcedony locule cast with remains of the endocarp adhering apically. UF 6297. PC, x 7.5. 
12, 13. Stereo-pair of a permineralized endocarp. USNM 424672. PC, x8. 
14. Same, rotated 120°, showing median line of septum. PC, х8. 
15. Same specimen as in fig. 10, apical view, showing tricarpellate construction, with one of the locule casts partially exposed 
(lower right). USNM 424671. PC, x8. 
16. Same as іп fig. 11, apical view, x 8. 
17. Transverse section showing remains of two of the septa, and large locules. USNM 424674. RL, x 15. 
18. Transverse section of permineralized specimen showing collapsed locules completely filled by elongate cells. USNM 435042. 


RL, 15. 


PLATE 61 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PLATE 62 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 191 


EXPLANATION OF PLATE 62 


Figure Page 

Ше: Tiffneyearpa seleroidea рей. et sp. NOV. Holop USNM 355045. е Ие 112 
1. Basal view of endocarp with mass of siltstone adhering, lower right. PC, x 3.5. 

. Stereo-pair, lateral view showing ellipsoidal outline, longitudinal fibrous grooves and oblique fracture, apex lost. PC, x 3.5. 

. Transverse section near base of fruit showing hollow central column and the base of three locules. RL, х4. 

. Transverse section at a higher level, intercepting seven locules. A line of loculicidal dehiscence extends from each locule. 
Three additional dehiscence lines indicate the position of abortive locules, so that the total number of carpels was ten. 
Etched, RL, х6. 

6. Transverse section near equator, showing central cycle of vascular traces alternating with the locules. Etched, RL, x 6. 
7. Detail of locule containing remains of seed coat, and showing polygonal cells of the endocarp and dehiscences line. TL, 
XIR 
SEIS. Dmparmsemenibönesirpen. et Sp оу. ш у у. ee О а о мы Л E уны о LM 113 
8-11. Silicified seed with seed coat stripped away showing casts of internal chambers. Holotype, UF 9288. PC, x 10. 
8. Basal view showing Y-shaped partition dividing seed into three chambers: two large lateral chambers and a wedge- 
shaped central chamber, each of these chambers represented by a smooth chalcedony cast. 
9. Ventral view, showing the hilum at base, lateral chambers and apical protrusion of the central chamber. 
10. Dorsal view showing full length of the three chambers. Note the facet near the base of the central chamber, and the 
longitudinal ribs at the junction of the septa with the (now missing) seed coat. 
11. Apical view, showing rounded dorsal side and V-shaped ventral side. 
12-13. Another specimen. UF 9296. PC, x 10. 
12. Basal view, showing hilar scar at the junction of the three chambers. 
13. Ventral view. 
14. Transverse section of a seed showing the Y-shaped septa and a portion of adhering seed coat, upper right. UF 9578. RL, 
x 10. 
15. Transverse section of a seed with embryo in central chamber. UF 9577. RL, x10. 


чл hh Y 


192 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 
\ 
t 
( 
EXPLANATION OF PLATE 63 
Figure Page 
1-1 2C ИИ О асове Sens GU SD: локва, aa о Ne 1065) 
1, 2. Stereo-pair, lateral view, prior to sectioning, showing two locule casts and remains of a disk-like structure near Ше apex. 
Holotype, UF 9472. PC, x 7.5. 
3, 4. Stereo-pair, same, basal view, showing two intact, broadly concave, smooth-surfaced locule casts; another locule cast inferred 
from symmetry (lower part) is missing. PC, x 7.5. 
5, 6. Stereo-pair, lateral view, prior to sectioning. USNM 354051. x7.5. 
7, 8. Stereo-pair, same, apical view, showing circular outline of disk-like structure, and trigonal symmetry. PC, x 7.5. 
9. Transverse section of a specimen in matrix, showing two of the three locules. UF 9480. RL, x5. 
10. Transverse section of the specimen in figs. 5-8, showing all three locules each with an intact seed. RL, х7.5. 
11. Transverse section of the specimen in figs. 1—4. RL, х7.5. 
12. Same, enlarged, showing anatomical details including layer of anticlinally elongate columnar cells and outer layer of paren- 
chymatous mesocarp. RL, x50. 
15-20 5 ei TESTA UT SATUS PODES ерте АЮ. atom deco Mese ut oae it, fi tere C M 114 
13. Fruit, basal view, showing large pedicel scar, three valves and rugulate surface. USNM 424798. PC, x6. 
14. Same, apical view, showing coarser rugulae and central scar. PC, x6. 
15. Larger, more triangular specimen, basal view. UF 5237. PC, x4.5. 
16. Same, lateral view. PC, x4.5. 
17. Small specimen, basal view, with two valves shed, upper side of photo, revealing smooth locule cast. Holotype, USNM 
424799. x8. 
18. Same, apical view, showing coarser rugulae. PC, x8. 
19. Same, lateral view. PC, x7. 
20. Specimen with two of the valves partially broken away showing thickness of wall. USNM 422396. PC, x4.3. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 63 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 64 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 193 


EXPLANATION OF PLATE 64 


Figure Page 
153107 Trisepiicarpium minutum пеп. et sp. noy. Holotype: USNM 24A73T ал nn 115 
1. Lateral view, showing pyriform outline, rounded apex and narrow, truncate base. PC, x12. 
2-3. Stereo-pair of the same, rotated 120°. Median groove occurs between adjacent carpels. Arrows indicate the levels of sections 
illustrated in figs. 5-9. PC, x12. 
. Apical view, showing circular outline and trigonal mark delineating the three carpels. 
. Transverse section showing thin fruit wall and thick septa dilineating three locules with flattened seeds. RL, x25. 
. Transverse section, showing flattened seeds within the locules. Acetate peel, TL, x25. 
. Transverse section at a higher level, with a large central triangular space and flattened locules. RL, x25. 
. Transverse section near the base. RL, x25. 
. Transverse section near the apex, with one of the locules broken away. Note axile bulges, suggesting subapicial axile 
placentation. RL, x25. 
10. Transverse section enlarged to show columnar cells of the septum and reticulate seed coat of the seed. Acetate peel, TL, 
x65. 
LIA runcatisemensanotoldes Вене ор mov QUEANT DENN UM JU I M 115 
11. Lateral view, showing smooth surface, elliptical outline and oblique truncation. Holotype, UF 6459. PC, x4. 
12, 13. Same, stereo-pair, ventral view, showing narrow width, and a central depression within the elliptical outline ofthe truncation. 
PC, x4. 
14. Lateral view of specimen showing thin seed coat partially flaked away, longitudinal crease that may represent groove in 
embryo or endosperm, raised hilar scar at center of the oblique truncation. USNM 424650. PC, x5.5. 


које о о тъ 


194 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 
А 
i 
| 
EXPLANATION ОЕ PLATE 65 
Figure Page 
ОО ОРИ ПО CS T Ob ром, о ee N. ое. 115 
1. Chalcedony locule cast, lateral view, with some of the endocarp adhering to the attachment region. USNM 414509. PC, 
25. 
2. Chalcedony locule cast showing concave scar of attachment, micropylar/stylar point and relatively smooth surface with fine 
punctae. UF 9566. PC, х2.5. 
3. Same, transverse view showing oblong, rounded outline, and complex attachment scar with numerous protruding vacular 
strands. х 2.5. 
4. Ѕресітеп in matrix with adhering endocarp, oriented transversely, showing complex attachment ѕсаг. Holotype, USNM 
355363. x3. 
5. Same specimen in same orientation, partially sectioned to show wall thickness. х 2.5. 
6. Same, detail of endocarp wall showing two lacunae or resin ducts. RL, x25. 
7. Endocarp wall showing thin uniseriate locule lining, surrounded by a layer of anticlinally aligned small cells that grade into 
an outer layer of larger cells. RL, x 40. 
8. Acetate peel section of the same. TL, x 50. 
ОТОО NSIRDOIet O EA ou UM E viis WU нн буз сузы tabes ons o CA ке КИДЕ D E ји: 116 


9,10 


. Stereo-pair of fruit prior to sectioning, with apical portion of pericarp broken away exposing the locule cast. PC, x4. 

. Transverse section, showing single locule with thick wall. RL, х5. 

. Same, more enlarged to show somewhat shriveled seed coat within the locule. Note dark layer of locule lining (endocarp) 
surrounded by anticlinally oriented cells of the mesocarp. RL, x 6.5. 

. Same, basal view showing nearly circular outline. PC, x4. 

. Same, apical view. PC, x4. 

. Detail of pericarp showing anticlinally oriented short fibers. RL, x 20. 

. Same, showing columnar cells of the locule lining, many containing rhomboidal crystals (seen in section as white squares). 
ERIS x70; 


PLATE 65 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 66 


Figure 
1-10. 


11-14. 


15, 16. 


ДИ aer 


CLARNO (ЕОСЕМЕ) FRUITS AND SEEDS: MANCHESTER 195 


EXPLANATION OF PLATE 66 


Carpolithus bellispermus Chandler. ..... esses eese eet eee hehehe hehehe ehe = 116 
1. Seed, dorsal view, showing ovate outline and intact asymmetrical hilar region. UF 8544. x16. 
2. Seed, dorsal view, with well-preserved seed coat, broken apically. UCMP 10696. x16. 
3. Same, ventral view. x 16. 
4. Complete specimen viewed ventrally with transmitted and reflected light showing silhouette of rounded endosperm cavity 
with obliquely truncate apex. UF 8545. x16. 
. Internal mold of seed coat, ventral view, showing low, rounded, median ridge and obliquely truncate hilar region. UF 
8554. PC, x16. 
Same, basal view, showing rounded dorsal surface, and low, rounded, median ventral ridge. SEM, x25. 
. Transverse section showing tuburculate seed coat. UF 6607. x20. 
Same, detail of of tuburculate cells comprising the seed coat. x 40. 
. Detail from fig. 3, showing tuberculate outer surface of seed coat. x35. 
10. Paradermal view of cells making up the inner layer of seed coat observed through the translucently preserved outer layer 
of seed coat of seed in fig. 4. UF 8545. x80. 
@ärpoliihüsspele USNM 353086 PE KI ОЕЕО nl O cro 117 
11, 12. Stereo-pair, showing elliptical outline and longitudinal ribs. 
13. Basal view, showing stellate central scar and radiating longitudinal ribs. 
14. Apical view, showing cleft, left side, and prominent longitudinal ribs. 
Garpolithus врео USNM ASATI m nennen ee т ашыны ee 117 
15. Achene, lateral view, showing truncate base, rounded margins and cratered surface. SEM, х20. 
16. Same, tilted to show base with smooth elliptical scar. SEM, х 20. 
Garpolithius spas USNM 3550600 to om. р с К От оү ы wen sr a 117 
17. Lateral view, showing wide elliptical outline and smooth surface. PC, х5. 
18. Transverse section, showing the thick wall of seed and pair of thick cotyledons. RL, х5. 


чл 


O o0 0 


196 


Figure 


1-5. 


13-16. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 67 


CODI SEA MIDS AAG 925 шарты N ee ae Е m7 
1, 2. Stereo-pair, ventral view, showing median longitudinal ridge. PC, x8. 
3, 4. Stereo-pair, lateral view, showing smooth surface. PC, х8. 
5. Transverse section, showing the seed coat pulling away from the locule at upper left. RL, х 15. 


p GIO sape S. WIEN Н marmot ee ои р кыл ee UE 117 


6. Transverse thin section, showing a single locule with a shriveled seed coat. RL, x 20. 

7. Same, enlarged, showing thick endocarp surrounded by a uniseriate layer of thin-walled rectangular cells. RL, x 50. 

8. Detail of fruit wall, showing interdigitating cells of the endocarp, the surrounding uniseriate layer of thin-walled rectangular 
cells, and an outer layer of smaller rectangular cells. RL, x 150. 


NY EI GIA RAS) O Ih Ni IN es dE TI о ДЕЛО IA WO е E е SIME. n му ыыр и LC IE CIT MN MEL TUS 118 


9. Fruit in lateral view prior to sectioning. PC, х7. 
10. Basal view, showing more or less circular outline, and central attachment scar. PC, х7. 
11. Equatorial transverse section, showing thick-walled seed or endocarp with a keel in plane of symmetry and a veruccate 
outer surface, surrounded by mesocarp composed of large cells. RL, x10. 
12. Transverse section at a level 1/3 of the distance from the equator to the base, showing placental attachment area. RL, 
x10. 
GOP IDE Asin МУЉА МОДИ i. ОО LIU көзун ШАЛЫ A Алде: МИ PA Hd IB RETE 118 
13, 14. Stereo-pair, apical view, showing smooth to wrinkled surface, roughly circular truncation. PC, х4. 
15, 16. Stereo-pair, lateral view, showing the domed apical truncation, rounded sides and base. PC, x4. 


| 
| 
| 
| 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 67 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


PLATE 68 


Figure 


1-4. 


5—9. 


10-14, 


15-19, 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 197 


EXPLANATION OF PLATE 68 


Page 
Garpohthusssp. S USNM 422526 а а он минава gin, fare eene е tere des lagen und бен 118 
1. Lateral view, showing truncate base, rounded margins. PC, x6.5. 
2. Basal view, showing circular scar. PC, x 6.5. 
3. Tranverse section showing thick wall, and single locule. RL, х8. 
4. Transverse thin section, showing wall made up of more or less isodiametric cells. RL, х 50. 
Carpolithus еро deu со сине о eet a ER за ура ee е eee MM MM 118 
5. Specimen viewed laterally, showing one of the three longitudinal keels, and protruding basal projection. UF 9556. SEM, 


x 14. 
6. Same, tilted to show triangular pattern at base. SEM, х 14. 
7. Specimen viewed basally, showing triangular pattern, central circular area. UF 6457. SEM, x 14. 
8. Apical view of a specimen in matrix with part of the outer wall intact. UF 9550. SEM, х 14. 
9. Same as fig. 7, light microscopy. UF 6457. RL, x14. 
Gürvolithus 5р. 100.2. te eed e же ис heed oc te i EI cee aria Ае sien, io ы te 119 

10. Ventral view, showing triangular outline and elliptical hilum/funicle. USNM 435102. SEM, x21. 

11. Same, oblique lateral view, showing depth of seed. SEM, х 23. 

12. Similar seed, ventral view, partially broken apically, showing curved embryo cavity within. USNM 435103. SEM, x24. 

13. Another specimen, apical view, showing keel (fractured) in line with hilum. USNM 435105. SEM, x23. 

14. Same specimen as in fig. 12, lateral view, showing relatively flat ventral margin, convex dorsal margin. SEM, x 20. 
Garpoliihusisp:- Lis USNM 424695 POP Он се еле 119 
15, 16. Stereo-pair, ventral view, showing lateral ruminations that are interrupted in the medial area. 

17, 18. Stereo-pair, dorsal view, showing median keel, and lateral ruminations. 

19. Lateral view, showing ventrally concave curvature and narrow outline. 


198 


Figure 
1—4. 


13—16. 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


EXPLANATION OF PLATE 69 


Page 
УУЛА Пазара ела СЪ КИРО a s ee А en ШТО 1019) 
1. Chalcedony cast of fruit, lateral view, showing smooth rounded surface, with meridional grooves between Ше carpels. PC, x 3. 
2. Same, basal view, showing five radially arranged carpels. PC, x3. 
3. Same, apical view, showing a median keel on each of the carpels suggesting loculicidal dehiscence. x3. 
4. Same specimen, sectioned transversely, showing radially arranged thin septa, some of which are ruptured, and empty locules. 
REY XS. 


SOFORT M oo Иш AACE RE ag DE Tc ceo И ИН Пн Па AMET 119 


5. Lateral view of endocarp with adhering bits of siltstone, showing ovoid shape. х3. 

6. Transverse section showing thick, more or less circular endocarp separated from the outermost thinner exocarp. RL, x5. 

7. Same, enlarged to show endocarp composed of short, anticlinally oriented thin-walled cells. Locule is at top of fig. RL, x 40. 
8. Same transverse section showing exocarp composed of densely packed sclereids. RL, x40. 


OA) DONT US: су ы VIRGO OU: cc PM M M x Mal папина НОЩ cas ate UE iile) 


9. Lateral view. x3. 
10. Basal view, showing central attachment scar and circular outline, x3. 
11. Transverse section, showing central group of seeds and surrounding parenchyma tissue, x3. 
12. Same, detail showing three pairs of seeds, one pair fully developed, x8. 
ОДН ЕВЕ WE Ола КА re E M GN NM IE Ee. IU UC ERN ET LEE Im 120 
13. Ventral view, showing concave surface and two-lobed morphology. PC, x8. 
14. Dorsal view, showing convex surface and apical and basal clefts. PC, x8. 
15. Lateral view, showing strongly curved outline. PC, x8. 
16. Ventral view, enlarged to show striated pattern of cells on the surface. P751 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 PLATE 69 


PLATE 70 


PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


Figure 


1-5. 


6-8. 


9—12. 


514. 


WS AG 


17: 
18. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 199 


EXPLANATION OF PLATE 70 


Garpolithusisp ДӨ er ee АЛ ee ee ла RE 120 
1-2. Stereo-pair of a block of sediment containing numerous seeds preserved as molds of the outer rugulate surface and casts 
of the inner seed cavity. UF 9794. x4. 
3. Internal mold of seed coat nested within the carbonaceous external mold of seed coat, viewed laterally. UF 9795. x 10. 
4. Similar view of another specimen. UF 9798. SEM, x 10. 
5. Silicone cast of the specimen in figs. 1, 2, showing rugulate surface pattern of the seeds. SEM, x 10. 
атор D ОК ОЧ Н. уууч onu е ааа лета O ae 120 
6. Locule cast with some of the pericarp adhering, viewed laterally. Equatorial cuts indicate position of transverse section 
shown in figs. 7, 8. x6. 
7. Transverse section, showing thick wall and locule with a single seed. RL, x 10. 
8. Same, showing parenchymatous layer surrounded by a uniseriate layer of anticlinally elongate sclereids. RL, x25. 
Garpoltthusisp voli ШЕ O49. и и ccs Т nee ee old asse UI а. O нв ono du Шы 120 
9. Basal view, showing compressed hexagonal outline, with central pedicel, and median keel. х 5. 
10, 11. Stereo-pair, lateral view, showing thick pedicel attachment and rounded apex. х5. 
12. Transverse section, showing thin fruit wall and empty locule. RL, x8. 
Gamol рО USNM A ЗОЗ АВО o ME Lc Mtt DUE oM EHE ENT ex 121 
13. Lateral view, showing rounded outline and longitudinal striations. 
14. Apical view, showing triradiate keel. 


Ganpolithussps20. ШЕОЛ ЗО e A e зл en ча мз йык шыга и 121 
15. Transverse section showing nearly circular locule lined with quartz crystals, and irregular thickness of the eroded pericarp. 
RL XS 
16. Same, showing detail of isodiametric sclereids composing the pericarp. RL, x60. 
Five paned Ноте Ие Ааа en E E орна САН 121 


GO) DOHTNUS:S Pe 21. USNM 422525, SIS се мрз O ОИ URE a ee ae A ees 121 


200 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


NOTES ADDED IN PROOF 


1. Recent geologic mapping in the type area of the Clarno Formation has provided new information and an alternative interpretation of 
the local stratigraphic succession (Bestland et al., 1994, and written communication). The porphyritic unit immediately northeast of the Nut 
Beds, previously identified as an andesitic plug (p. 9), is now considered to be a dacite dome, composed of plagioclase-hornblende porphyritic 
dacite. Bestland et al. conclude that this dome was already a weathering topographic feature at the time of Nut Beds deposition because bake 
zones and alterations of the surrounding sediments are not evident, and because reworked clasts of the dacite occur in conglomerates within 
the Nut Beds. Whereas C. B. Hanson (1973) interpreted the Nut Beds to be separated by an unconformity from the mudflow deposits of 
Hancock Canyon, Bestland et al. consider these units to be approximately coeval. 


Bestland, E. A., Retallack, G., and Fremd, T. 
1994. Geology and age assessment of late Eocene fossil localities in the Clarno Unit, John Day Fossil Beds National Monument, central 
Oregon. Geological Society of America, Rocky Mountain Section, Abstracts with Programs, vol. 5, p. 4. 


2. A samara of Cedrelospermum was recently recovered from Messel, Germany, confirming the presence of this genus in the Middle Eocene 
of central Europe. In conformity with other European representatives ofthe genus, it has a terminal stigmatic cleft and lacks the small secondary 
wing typically found in North American species (Manchester, 1987b). The specimen, Me 7274, is in the paleobotanical collection of the 
Senckenberg Museum, Frankfurt and was examined through the courtesy of Prof. Friedemann Schaarschmidt. 


3. Fruits of Cruciptera have now been described and illustrated in detail on the basis of specimens with excellent cuticular preservation 
from the Middle Eocene of Messel, Germany. Cruciptera schaarschmidtii samaras are about half the size of C. simsonii samaras, but the two 
species are difficult to distinguish morphologically. Epifluorescence microscopy of the Messel specimens reveals that Cruciptera possessed 
peltate glands, another character attesting to its affinity with the Juglandaceae (Manchester et al., 1994). 


Manchester, S. R., Collinson, M. E. and Goth, K. 
1994. Fruits of the Juglandaceae from the Eocene of Messel, Germany, and implications for early Tertiary phytogeographic exchange between 
Europe and western North America. International Journal of Plant Sciences, vol. 155 (in press). 


4. In addition to vitaceous seeds, the grape family is represented in the Nut Beds by anatomically preserved wood. Based on a thorough 
survey of available extant vitaceous woods, Wheeler and La Pasha (1984) recognize two distinct taxa in the Nut Beds, both of which are 
assigned to a new extinct genus, Vitaceoxylon Wheeler et LaPasha. Although the fossil species are vines with anatomical characters diagnostic 
of Vitaceae, they do not conform to any one extant genus of the family. This new information from woods raises the intriguing possibility 
that seeds morphologically identical to those of certain modern genera might have occurred on vines that were anatomically distinct from 
modern genera. 


Wheeler, E. A. and LaPasha, C. A. 
1994. Woods of the Vitaceae—fossil and modern. Review of Palaeobotany and Palynology, vol. 80, pp. 175-207. 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 201 


INDEX 


Note: Page numbers are in light face type; the page numbers for principal discussions or descriptions are in italic type. Plate numbers are in 


bold face type. 


ND DES RTOS e DRM hea UTR ee 30 
Аааа ian re M UTE 85 
COIN Marre ым АА БЫА Ыыы Eee Yirg 47 
COSA FLA Oe e du naL MM 2i 
ЕЕ ОЛ ЛЕ ОЕ Ж. E КЫ р аы A 20,27,33-34 
MATA ASUS AAN a Аа Е тр 4,16,33-34 
arcaica ee 33 
A A E 34 
AU ee yee 20,27,34-35 
САМИ Зори TOV ER VER ae CD rS кус КҮ с АНШЫ: 4,16,34–35 
GresOnensisuscotmand, Wheeler ce ЕИ 34 
КОКС ТОМА ОМА О ИЕ 4,16,35 
Gao orar te RR UNE 10 
Па аре oor НИНА ta ee PAR ORUM 24 
Оаа и ОКО eit e не TO ES 20, 27,93–94 
ОШОО р MOVE whe se ha pe aba ala 44,17,93-94 
SIMA A his GRE SER 44,17,94 
AA A ee 20,27, 94-95 
ROOSEALSDNON ET АН lis EN 44,17,94-95 
Anacardiaceae Eindley c o 5 и а 35—37 
ТИ ЙИ СО] Ө нал cies D уз уу EAE REN 20,27,66 
CAS аны c ылла ЫК И ae КН 27,66 
(С СОИ loh кома ара трак ова о 27,16,66 
ОРО У еј ONA ee a 96—97 
VALIA MN Улу EE RC CUI р О 46,17,97 
nOs ane ыу ы ee кй eee a 37-38 
Anonaspermum Ball emend Reid et Chandler............. 37-38 
OESS DIO у ллу a лу К ee N 6,16,38 
cf. pulchrum Reid et Chandler ................... 6,16,37—38 
ОИИБ ПИО Vas ТЕ Du uo Md ла ZO 
ООСО ОЕ 97 
ОРОЙ У MONG ет оа IA 45,17,97 
ОВП НОРА ТОНОВА eu 20,27,90-91 
МИК] TEN OSA. MINUS ee US Ot 42,17,90—91 
ЧЧ аш Ms vo taie re decre re eos AR 24 
A ER а ате 38-39 
A ОСА От 25—27 
Боасе помела ee оа 97 
ATA NO 46,17,97 
zAsplenium" hurleyensis Berry .. узш. 20... кошу тА 2i 
OAI EE е лу уул сушка» ee 71 
СТАРИОТ ee 30,16,71 
venablesi Chandler) Chandler sc ee 71 
AUO AAU A MOV A ууну сл суу Uo MIS UE 97 
СПОРОВЕ А РОМ vee ocio ae ae cro Пеле, 46,17,97 
Danamaspmilyi uten ER 73,15 
ЕО И ERRARE LA ic Cn 114 
Bound oou ААК eee ee te эк Coe E 25 
АО ee S ыча. сс мс 20 
Bava Ia Оууу. a aN a I DR EE 39-40 
(око еден ср ВИ nt зунда Dee LO 21 
ШИ ЧЕ ык к аыл ea NENNE et 
E ICH OIG ee ee UE RS 7 8,38,82,98,105 
Bonnia GENE MO Veici ienen ae, RES APA АА 98 
сраста PEDON оо RN k DUM 47,17,98 


Brandon lignite, Vermont эс нул ee НЫШ 24,35,65,66,89 


BrummensSpruneslorar а соса a me Ж 10,13,44,59 
ЖОШО а S ieee ae A е ао ы 20,27,86-87 
IDO ESO ee 39,17,86-87 
SUDAR CUM о зао РАЛИ хым җы: EL a 39,17,87 
Burence И gti Mert се aa ic tilt ten O 4041 
ВИЕ Reid et Chandler nr en ие 40-41 
BIAWICKERSeS@handler а ee en 41 
OPE CONCHE SOE NON слан у шугыл ee e EE 8,16,40—41 
ee ы Са ide ee ter 8,16,41 
Calycocarpum Nutt. ех Ton et Gray... 20,27,69-70 
AE ба ИВО ТУЛ пива ak say ei Gis a 28,29, 16,69-70 
ШОШ (Пи еп К Gat Ду en 70 
CODO RUSSO Ое nn... nern 15,28,117—121 
ЕОР Chandler. en a... ВОЛАНИ ee 66,117 
ри Reid ei. Chandler) Chandler... с с ли er ae 89 
О ee ЖА ЫЫ S сы 66,117 
A ИТ ы т т 66,117 
du ege Ce aer MeL VE MU ME ES e 66,117 
ОА уо a d A са пое зли у у 67,117 
TA E Иа 67,117-118 
d OMS MERE EMEN RI EM IM. с Eo b 67,118 
з, ЛЕ Ей os A QUNM И IM ра EE o ва 67,118 
cobi E MT ME S ccc S у; 68,118 
Sire Ar е ү аа CMM DE EE 68,118 
SPA mM MEAE ED pa A RE 68,119 
corde AR EM MMC ru LO 68,119 
STOO иеа ee Ce И ARE ПАЛИ 69,119 
SDR Se de I CL Qu Ur. то ва ru ER ANS NS 69,119 
den dod aeree A ы; ЈУ 69,119-120 
сү а y AMONT PEUT alls се 69,120 
dou Mac m irene бу E Р ARG cane ame 70,120 
PES da лыо и een Sea Уны а Лаб AMD 70,120 
о Й de Me Кыс ын Ыы M 70,120—121 
Кр И eph кае е A А УЕ 70,121 
Sese eU ee een Er 70,121 
СБР НЕ еа Е нк 70,121 
GUSTIROPSISAO. DOES раси о о ай Jet 20,27,46-47 
Ларе SDN e ah. AA Ren en 12,16,46-47 
ИШОНРО TANCK ее. (Саша. 12,47 
CCANEIOSPEV SOPOTE Пара doses das Ine ees 25,92,200 
lineatum (Lesquereux) Manchester .............. 43,17,92-93 
СНО МОЛ Жо ен MERE N 90-92 
acu c у оз ee NN ON 20,27,91-92 
DAA у а те Н ое ИЕ 42,17,91-92 
SI ee ee Tt WER + 43,92 
CST PAINE Са a КЕЛА eee essen 26 
AUS CS ee ee RS 66—67 
TOC MN OSQESGOUDE у седла хуу ae Doe d 27,16,66—67 
КОШУ Ci AO A a 10,13 
(EN лош: yore reid cin. A A SON 24 
(CANA a A PON л 8-13,200 
“А ОЛАЙ ШЕЕ ee сы. „ОЛДА. ДЕЙ 78 
Clarnoxylon Manchester and Wheeler ................... 20,57 
мА сйс 1 ооо ie Oa ai reel ај QU S 20,27,89-90 
Теа о е НИМ 42,17,89-90 
Сота A ee ИНИ 66 


202 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


а TREN, MOVE Sere ee Не 25 
HUN HDD ол EM а EUN ah 19,16,55 
(СОРТИ ИТ A Sen OVES S IE a LUE 98 
CHICO sio d er В Е a a 48,17,98 
oa ОО OUO TCR TUNE Arv dus dete 41—46 
En Huit ГАРОНА 9983 20,27,41-42 
CI HOAs RE ен, IER 9,16,41-42 
Са О ee Dove On ЬО И 39-40 
e кипи ONE A a ооа 7,17,39-40 
A A А REIN 40 
QUOD NC SUE е WIE Evans. sisse у корея ухо nee 18 
Ci eU UM a ir Leu M es due di A 2l 
СОСЕДА ОПЕНА и kc cial ern 57-58 ,200 
ПИО (Brown) Manchester „au... 21,16,57-60 
COIE Ms OVI pet Is ts ee SAIN: seek UR ut He 98-99 
И Со по и Wt cu. GCE A 47,17,98-99 
СРО ОЛЕ реш ШОМ En 71—72 
MOO eS We Е ТО СС ЛЕ мана Joh MEA 31,16,71-72 
ОУ ee mean m ди o ОА oe 21,23 
DAVIS COPIE СПАЙС пасти u er au му Лл etit уе; 73 
ОООО СВОД ЕТ О а ЖДО. жу аи ИЗ 
ПРОСТОР БАВИТИ СИ cer AAA ДЫЙ. n3 E 31,16,73 
DEE CHE Yos Rack Dona куык eo ERU en 20,27,63-64 
ЈЕО ОРИ vau Bea eg een el 25,16,63-64 
ү ы ро DU DM AU ad c E N 21 
A a a ые Ер ee 99 
PORTO iia me ee C ERES 46,17,99 
a A ION da de RR TNNT 85-86 
ОАО кык T un 39,17,85-86 
НО ea ES ааа 21 
ЭШ ЖИЛ MINGUS: съда ра дъ а ew cea he es 16,20,27,73-74 
auriformis (Hollick) comb. поу,.................. 30,16,73—74 
PUNO RE E ae скы буа ee ot 74 
DO ODUM аре land Mazze c. cer ee ner 18,41,44 
PUD ODO ee ат nn LE РЕС. 31-33 
TOPES SIE TON иа ia ри hun we лаје 3,16,32-33 
РЕВЕНЮ У ne a ER 21 
ШИП Roid et Вин злата слой iu error IN A 42 
y EUA БОП. ШШ аи кик A ЕСЕ, 99 
ОАЕ жы он А ы и А RU ard рова 48,17,99 
О ИШ ИШЕ ШИ ә», en c «tuer etes пина Ss cE e 28:56 
ПАО Кој Па а GAC NG cos НЕ а en ce ај ee м 10 
Fasten Аай, шаш Mora cuoc ватра Л, IR 25-27 
easterm North America, extant Пота... одн 25-26 
aida DEV. A E aT 20,27,80-81 
a A A т тг 36,17,80-81 
TOV Ухх Sata ko s oC UR bd. EIER 36,81 
ВИ ION AID КЕШЕШ uec cem y eque thee hs y CX OMA 59-60 
ЖОШ Nene ER TEMPLES EE 59-60 
Engelharhoxylon Manchester... bused csv con ка ted 20,57 
PMS CTC Ue eu Me Season ec de жу оку ЕККУ И 20,27,75-76 
oregonerse Manchester et Kress...............s. 32,16,75-76 
BolypserraBerdiet Chandler o ыу алуын 74 
Pornonun Reid а Chandler... me 74 
И о A eue kx d OH en 29,16,74 
WITNESS ЕА uer eere ууу»... и 1510575521 
ПИО Ve SUOW O NEED Les 15 
ЕШР ne AMMO so ve e erre edges АО Ар, 26 
Bunope, tossilktlora. 0... miese Аи 8,24-26 
Pocs I=3 0f the Nut Beds....... ee ee Pier? 
ЈРАВИСОНО IIE а So evs vs sin pen We wee И Oh M 46-47 


СИА MONS COLS ENVASE nn 20,46 
ТОРО cA ни сыы Ет AER NI M Cree On Mee S EI varied TELIDA 
BRERFIZNOCANDUSF T NO TTE ee, RA UA ЕО Re 99—100 

DIVA INSP AN OV aria MO MUI ER Lt SET 49,17,99-100 
да ANE допира ON NA ent Cee в аве ае 100 

WINES DANONE а ДИН ToS Aa m 50,17,100 
УСЕВА ОМОТА со ода RE бати 70,121 
Еавошвнасвав DEAN ИТЕ О, ТРИ ER 47-49 
элне ж е Кк жү ea es IE 15,18,19 
ЖООЛАП ES E ШОУ ar E ies Pik A S eres 49—50 

ОЙ CAST Soy AMON УУ о аа Wey ee сър a 14,16,49-50 
TARA X S UR ONE о NI 100—101 

ЖЛ ЛИЛ ЕРГӘ ХУ хы ыкка лт ELS 50,17,100—101 
Пена КЕ отат ре msn. en dete күз на ш 12 
(69 бат етан кобалт Г. 24-26,77,84,88,89 
ПСО: О eate me Ho WF Bart озы 15519 
(А Сир IT BOM. NOS УЛУК К ККУ Г К КК 101 

LAMAS MN О СОЕ ТИГУ 50,17,1101 
Green River Formation Ora r rers 49,76,93,94,95 
ЭШЕ е иеа 19-21 
Hamamelidaceae КИВ оо И Ий 49-50 
асос Canyon ne A O ON 9,10 
FIA OOCKM@UALLV ates Een etu КӨЕ ЭНДО. 15 
NEL E O дер лой ы АУА O S, 21 
ере ОСА о M а ee 101–102 

EME AA RA ва et УА 51,17,101—102 
ШЕОЛ Ore tice tev A үис ви RE ER 10 
Ж ИЛЛ Л КОЙ ЫАЛ ee ara 20,27,50—51 

GOTHIC SD нл н каз жеу ж киж ул а OU „ҮМ РА ИМ 16,51 

КОЛО HORUS D RMON ee 15,16,16,50-51 
Тада ЕТ отот У кр ну Fco 50—51 
PAVO UD US ы N ЫА ee RD 119221] 
Itaca Mid EE КАНЕ 51-57 
Пе а КОНА 1 eure о Ау OS 96–121 
ШЕШШ КОСЕ ине TN В РЕТ 10 
СЕСЕ а CTA НАЙ 18 
IU de ne A 51255 
ООУ БИШЕ IE e IOCIS Le Rr uis 20,27,51-53 

СИОН ВУНОМ I ee 17,16,53 

COSA ne ie ы А а Ж 19,52 

multireticulata Reid et Chandler ................ 17,16,51-53 
ТООДО ДЮ ОУ pa Г MET И 53—55 

ООА Вр ПО а SEMI ee 18,16,53-54 

lentos Spo e t ада OO AN 18,16,54 
UCC Ones ROOM ONT Кызны ое 102 

OO ШОШ КО, уул ы УУ ЛУ С ЛЫ 51,17,102 
John Day Fossil Beds National Monument..................- 9 
Ж ШЕШ К ОШ Ший у. ер и и НА ИЕ НИ оу л 10 
Juglandaceae A. Rich ex Kunth MUR UR A Ыы сы е NE 57-60 
Шш e cC nM M E СИ 57-59 
OGIO qup M E M EE ус 20,27,58-59 

оО оо 21,16,58-59 
ETSI DEMISSA en ee ae Ае 40,88 
ЖКО ОЛА ОЛ кози o a ario RUN 40 

CI LA те Meer E ET 8,17,40 
КҮШ Formation, Alaska... е 24 
Ооа Mite Ота со nen жуы rt NS 12 


БИКА Formation, Alaska. Do Пен лыкты лел 24 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 203 


Haro ma Reidiet Chandler е У 44-45 
Саа Reid et Chandler erh 11,16,44—45 
Парот Ога не DOR RIEN Ыкы n MENS TENE 29 
E N 60—63 
плови Reid et Chandler eme лз ОЕ 61—62 
СЕТЕР РИМА OW ash А 23,17,61-62 
Laurocarpum Reid et Chandler ......................... 62-63 
VO SOUL РОМА NT RA RERO Пи И EAN 24,16,62 
обор Е но БИО S c SEIT 24,16,62 
тако а Rede Chandler a sn ae 62 
CUNAS оно кора SUNT ERE TS NEN 24,16,62–63 
Пе И а о A E un 8,9,12,15,18,22 
Шеонилосатрот Goeppert orenen E a 63 
SDI sd nO S S AS DEM 25,16,63 
osa ее d DN 63—64 
ШИА ne ee Rm кисы М 19-22 
И AS E N OVA ee 102 
А S IN у E E E ED ee 51,17,102 
ООРОО 102-103 
SOS CIRO ОСН uo Le EN eig 53,17,102-103 
O tL Каты ME 24,73 
ШУ О ТАЛКУ ee ee О о ох оо 20,27,61 
GIO HOSISESDU LO VoU a CU IER TD 23,16,61 
london lav Horde t A NN ON CUM 
DER 8,24-27,35-38,41-43,45,48,49,52,53,55,56,62,64- 
66,70,71,73-75,77,84,86,89,95,96 

С ӨЕ ДОУ хол TU TU с 103 
CURVAS DADO ое 82.17.1705: 
ШООЛА URW) M 2i 
Троа OS ЕНИ УУ ы tao RS OUT erbe 63-64 
И ойра Manchester ee 77-79 
sabta Manchet a пера Ау 33,16,77—78 
Macginistemon mikanoides (MacGinitie) Manchester ......... 78 
ШИ ср о Manchester i у. у л ТАКАЛ ОРТ 77-78 
ОРИ ИТАН Лл c RE rd ele Lead a raum 20,27,64-66 
ОООО УЛОУ NES УИ 26,16,64—65 
PAO ОШ ИА ANON ОЕ К Ул EDI UC И: 26,16,65 
ППИ А ADAMO ата ЕЯ 25,26,16,65—66 
Му Оа саб“ Бы Уло у ыу. MAN 64—66 
ашан Ел CO CONTRE с АЕ ee 9,12,13,21 
MOS Bl те О EO 20,27,42-43 
он ын leto aie LA EOM Па RM uA dr cp np Ee Pee 10,16,42-43 
INI COT DI @handl er eine о аак 44 
NLC AME Ще Соран ON A Sarno cut Кл УМ es aN E a л net 10,44-44 
ООА ТОСОО SCOUW О Е 43-44 
ОРОО у SCOUT е 10,16,43-44 
O Ma BIS I TREE IN 20,27,81-84 
ле е IMIS ORM ОН E D CU DN 37,38,16,82 
WOME о од ee 37,38,16,82 
а ет ди ООУ ан aia з E 37,16,82–83 
аллома DON А ил Ы 37,38 ,16,83–84 
а ува а eM аСТ СОИ NEM 37,38,16,84 
DdSnispomHacedcM dS NER а RE 66-75 
Маон A E у ХОК av: VS 
Меза (О јавите њи а OTT TS 24-26,53,55,58,118,200 
TOS те teles НЕА деу 18,24 
ШО ОО ИК ЕШ ПОЛ en 103 
ШОО КИЕ ШОУ CI RUIT DUE кзз чаж олжосу лон» 53,16,103 
ОИ a Вас er ees 75—76 
Мо и M MO LL а e 75-76 


(ООО ОШЕН SCHE MON en en en PUB 103-104 
ПОПА ИУ Sell ONE а Е E 54,17,103-104 
Norm Amonoa. extant ота аа QU ER 25-26 
O A een A ee 25 
IN DACACEACS IN EL du M EE EMEN 21 
AA EB eH е а 20,45-46 
ЈУ сай BI RW RON EUR жузж a ТЫ 46 
AR Ware E Ce fe cance tein Ur MM MESE 12,16,46 
ЄР КО DU AL ек мА Н 12,16,46 
крайаей бой) combenoy. л 11,16,45—46 
falamanconauaamimel et Zamora. ee 45 
QUOTTOCATVOIGEH SOCOM ОЕ Seed eee 67-69 
ЙОЛ ОХ SEOUL mica ro Mince E EN EUER 28,16,67-69 
OMIC U CU MOVE er 104 
ФИН ES оь зоо A e e D SS 52,17,104 
ANTAD A A x E LU ali Dad dee 59,60 
ОТООР A Eee A S E 12.21 
Balneoallophrius Reid et Chandler a er aa 86 
ОООО nee le dl 38,17,86 
COUNT DANOS rs Bi oc I I елей 38,17,86 
PUOC OV чарок а we по у ck ee ee 59-60 
cf. Palaeocarya clarnensis Manchester ........... 22,16,59-60 
аео ууа епа е ann 18,45-46 
РЕ ёр лете Kad er Chandler gi... eu. da 55—57 
ОСОО епа еј Chandler nn кк Е 56 
ОВО И ЗВОНО ТАКАО ee 20,16,56 
Psevaopersica ЗОО omend. nn... 20,16,56-57 
Па О А опао |Кен ние н. 74–75 
PEDES јао и О er Ри Лл ужы E 30,16,74-75 
RHIC OCHINALE Hat had areas ee Me 21-23 
IPOlCODAnds Belle РОМ с cris coser eee и 38-39 
ER ORENSE MON Ere evt rd eg ah 7,16,38-39 
JO DIG GUT) Майонез. бр а RR ROS 60 
Paleopianyedryar hickepi Spe MOV оа ае 22,16,60 
Paleorubiaceophyllum Roth et Dilcher...........: 5 5 a 81 
АОВ O а DIE 76—77 
ООВ Eee Cn A ый 71 
BALNENOGISSUS Planchons ас a а a 20,95 
Quem ИСО СОБЕ ~ van are alba en 45,17,95 
ГЩ Ада К ШЧ ee le ee 45,17,95 
КОКТО UE SD ОЖ. E си онаш ducit: ee 104 
ОО ИДА ра а ен es ae 55,17,104 
PANA E Ine CNET ИН ИЕ Е 21 
АНС ВИСИНА РОТ NON. en ea ec vations ar 35-36 
minimus (Reid et Chandler- comb: HOY. .....uuu es 5,16,36 
Bhstoctene уы с ses en een 19,55 
Т туе ао еа О t eS ne 55-57 
ИЕ ОВАА ВА ио зае Bug Te мв 104—105 
FAS MOV и ОЕ ТТ 55,17,104—105 
О рее У COPI La Ace «елый DEL Mat 55,12, 405 
MAGS Аво a Co UE ra qu a AUS Um d. 30 
АД EN o eMe vba SOOO CLA UE celo еы 20,27,30 
IS SD ene ee cU Si DO ese cU rr cM ЧЕ 1,16,20,30 
IATA о aH UE И 33405 
IA O д. ET 54,105 
ПР ето ак о o Е ua 77-79 
TURCO HUS synandrus Manchester ..... da a 78 
ат Unsgeremendspaser. (а 9. 0... о 20,77 
TM СО а 20,27,77-79 
Палада а (е его (ода ee 33,34,16,78 
ГОУ УД ее ШШЕ ee ad: 60 


204 PALAEONTOGRAPHICA AMERICANA, NUMBER 58 


ПЕЕ E м eet een Bea ЕЕ СОТА 60 
A репе TION. ne УМИРЕ ла 105 
“О ШИЮ ИР ШЕК ео полне SEA: ЗЕТ. ПОК 56,17,105 
ite UI ON ea м.м, ee ЖИМ ТБ 106 
ИКОН SO E O 56,17,106 
Прити Кее Chandler... ae 114 
DSTI a rn ea LE и 13 
ТИШЕ VOM eM ШОТА ue nee 23-24 ,64,80,94,111 
O DESSEN NON е an na ee een 106 
с с ү он ү en. RT 56,57,17,106 
d NI M Ue a Бу E И 20,27,79-80 
GSO SD едК knee па ела 35,17,79-80 
ОО ВИ ТОУ OR кы a Wee oro aan Ge Кале 35,17,80 
a MICE DOE IU. suene deles ene Wow а уж. Бе ел 106-107 
DEDE BV Lu er ee 57,17,106—107 
Pogot Grip, Washington 2. ceo о ne 12 
КОО ШОРО ФЕН INO nen veo poste RD ed 107 
КОЛИ сл HOW en Се а ан ЈЕ onem exis кыз» 57.17.107 
ТОПО та TAD OMEN: , 2v, ne een 205021597 
TENA ae а онаи ET 19,16,57 
zenkeri (8р. Моше) Еке... nn TY 0D ОЛУРА 19,57 
ENEE ОБЕ УЫ А ee ЛЫ MN 107 
DCT OLE SN MON. ers ee уузу» р; Re 87.17 107 
ШОР UI ОВ эми CREME DEN TH чака tr МЕ 20,46 
1 SUB acd PR EC uA OR Le vtae EA Ue АРЫ Ыы реа пелети FON 20,27,47 
РОО РЕ MO ESTEE eL. а eda eremo rare nee ont 12,17,47 
идеи отта ЕО ОСОКЕ в 107-108 
ТО ORIEL ШИМ eed sete ee re ee 57,17,108 
ПСО эй ы-н o на О ee ге с, 12,13 
ЖОО ШШ ЛӘН ee Los A кык. ТО 12 
Republic Йога, Washington 
А а ут. к-к е д» кк җа ыы a АКЕ 20,27,36—37 
ПОПОВО E ЕО is sr rr eec AN 6,16,36–37 
Rosdewde USE, а ка o a eb Seinen ween ans 79-80 
ПАСВА USB оо Ер E ии йин E 80-81 
OD МИЙ uario A а Sak bae 19,20,24,27,76—77 
bracknellense (Chandler) Mai 00 21% 33,16,76-77 
Jenkinsii (Reid et Chandler) comb. nov............... 33,16,77 
SARI СЕН O ИШҮ CHA 27,84 
prefoetida (Becker) COMM: MOV. co.cc ssn EN 38,84 
арос ВЕ т eae eee ev RR RIGEN 81-84 
МУДРИ ИРАН ИТ САО ЕР NOV ick ata li mec eras суык REN, 108 
TEU РОТ NON неза 58,17,108 
Sape quede JUS xor ocu a re ee 84—86 
а АИ A ДЕН 86-87 
У ОДО ретт ит Reid et Chandler... И. 48-49 
О ИЛА БА Та et Chamdler-.. L7... TE ULTRIES 48-49 
CNO SOMONE are oi penn ceive I IRURE 13,17,48-49 
SCA CCA И BEN. NOV. nenne 108-109 
С ПО Вера lO VW ao d ER rk rive enous ee 59,17,108-109 
КО К sa поли ee 109 
ООО е а ена rennen 58,17,109 
Ч DOV oa О ем 109-110 
КОО ПОП ЫН ШОУ. en en 59,17,109-110 
УЛО ШЕШ een 20,27,87-88 
CEBO SSI D; DON та ИЛИ ROSA 40,17,87-88 
зсанзапаШасейе Bl, u... da ЙА ОИ LE SO 87—88 
пас риска Creek Попа NE M E 10 
тшу куук се an се, THAT 19-20 


ЗЛ ДЕЛО та СЕВО она Keen 110 
ESTAS DION er nn ee ҮЛ. НЗ 58,17,110 
УДЕСНО СНА ВОН MOVs ee a 110-111 
ООО ОТЕЛЛО UE ee TAT ы: 60,17,110-111 
Stapuyleaceae шау жй ти ~ oy Дена сол 88—89 
ОША ДЕ па и и Л лер Суулук кн» T тая 15,20 
BICUCOlY De: cons ASMO 2... were ee 18 
MOLLINA AN CORDE ee 111 
OOA орла ом е AA 60,17,1171 
ASU LAUSD EVITA Сепо е UA 111—112 
CORONGIU GLATTLAS ON на m TCR 60,17,111—112 
SI ЫЫ NR ууу кау DU ENT 28 
SyHplocacearrDestn о Un 89 
AS NEAL E TEA ee 20,89 
ПООРЕООО AS ВАЛО ууну». Ue C BET Динан 41,17,89 
DO ODIO СООО У 78-79 
СТОИЛО ера жил» бк IR 34,16,78—79 
КАО ЕМ КИНЕ Bean den з кз cate ЛЫ 20,35 
Тера сее е КОЛИМА ee 20,27,88-89 
ООЛО УУ ЛБ Ко гл уу кул Лы MAE и 41,17,88-89 
Ира те КЕ ara ee a nl 30-33 
TA A ee re 20.280 
дао aS ipl Ow Бола ууу a DIG 
ШООК САС ТОТА Мин ни Hodie E ced Mec ы ои 10 
TASA ОМ ЕЛП Уууу лз кс TL 44 
ШЕ ОЛ ЛЛУ A ете оло оъ о сы 122 
Теа МИА ПЕТОМ ОО ОСМЕ ИЛЬЕ 112 
MUDA NE о ә кит ош LM 61,17,112 
ТОЛ Л АЛИШ SVE ON ес ужы аза му мети и ни ie IN 79 
СОАТ ON уику Аук. узе лты ота же э 31,16,75 
КӨЙ ОА PDF EN ne IM 89-90 
Thelypteris iddingsii (Knowlton) MacGinitie................. 21 
ОО ОШКО ONS жы узук» eis) ко ro ld шыт te 112 
О СКОТ COS ANON E RO OU а 62,17,112 
Tinomiscoidea Reid et Chandler ........................... [1 
Tmo cden oceidenialis SP. Nov... 30,17,71 
OS DON CANES A к ЛЛ. улоу ы улке си: 20,27,70-71 
BIO SCHONSDENON RA etes s ik 29,17,70 
АДА UIA CASANOVA: DS TOU OC ER ER 29,17,70 
WIRING ISLC TCR fer Re OUT DETUR ж Л. NACE с 66-73 
ONE Qe MMU RENT EI OTI Ea у D Ted ОЕШ 
ЕЕЕ 2,16,31 
ER A RA ee 19-20 
ШО о О ук Eon ыда ЗТЯ 20,27,92 
PL GHG TONG [SEIN ON Nin о er КЫ Шыг 43,17,92 
VARIA ДӘП, МОУ... г» суму». Bagh oec arc ар 112-119 
OVER ONES SDENOV и Mes RR ee 61,17,112-113 
ЛУУ ANDI HALES ANNA ДД, ан а LIE ша 113 
ОЛОИШ ee, 62,17,113 
IDA ON Tate na 113–114 
COUME ONAE BD. ЛО о ba {уугу л йы и 63,17,113—114 
до евайце Ана дела (оди ЕК 114 
ОООО ае е 63,17, iM 
ШОХ ОИ ПИТ ОУ TURO IULIAE 114 
И О ба пета з А 64,17,115 
ПО А И О T NONE ne co cM cM куи ee Каа 10) 
ООЛО О COE к ур qae cct O aha аи ае MEC 64,17,115 
SURI ERW RI НЕ E c PEN Улу кй к кышкы СОГ. 25 
ШАА ИБО a er nn en an ee 90-93 


UAT HIE УЗА Л ЛО та RD tandem 20,60—61 


| 
| 


CLARNO (EOCENE) FRUITS AND SEEDS: MANCHESTER 205 


ОЕ es a при пита eee М 92-93 
Ulospermum ONE TL Vie Л а Te уж Ase: UR ut trae us MO у 

ШОО оом Уолл. T T 65,17,115—116 
КОР о и роуа URS ER 22 
CULE DTAte LOSSIS SECO T ASIN ya и а Е 9,12,13,21 
ђака а a cuc aL AME ен 19-22 
MA азса е 93-96,200 
Ne 20,27,95 


maenispenmasGhandle en. 45,17,95—96 
DU CNUs Dis ОМАН уу ao ена чн Ed 44,17,96 
UE, Ла ТОКИО ЕОНИ не 
ne 10,12,13,18,30,39,40,43,51,58,60,63,78,79,85,90,93 
аА er nee 9,14,59 
EA CR CLONE ONERE RR сан 116 
ПЕТА E EI ОМ ~ t tete Шык Ко c а 65,17,116 
И ҖЕ ОШ See ce прасе аи 10,13,20,39,40,51,58,60,78,79,85,90 
аа Трес ра e на лена ОЛЕ о E 21 
WOO шуо 8,9,11,12,15,19,20,22,34,35,57,60—61,63,77,79,81 


PREPARATION ОЕ MANUSCRIPTS 


Palaeontographica Americana currently appears irregularly, on an average of 
about one issue each year. This series is a publication outlet for significant longer 
paleontological monographs (i.e., more than approx. 200 printed pages) for which 
high quality photographic illustrations and the large quarto format are required. 


Manuscripts submitted for publication in this monograph series must be 
typewritten, and double-spaced throughout (including direct quotations and ref- 
erences). All manuscripts should contain a table of contents, lists of text-figures 
and (or) tables, and a short, informative abstract that includes names of all new 
taxa. Format should follow that of recent numbers in the series. All measurements 
must be stated in the metric system, alone or in addition to the English system 
equivalent. The maximum dimensions for photographic plates are 178 mm x 
229 mm (7" x 9"; outlined on this page). Single-page text-figures should be drafted 
for reproduction as single column (82 mm; ЗМ”) or full page (178 mm; 7") width, 
but arrangements can be made to publish text-figures that must be larger. Any 
lettering in illustrations should follow the recommendations of Collinson (1962). 


Authors must provide three (3) copies of the text and accompanying illus- 
trative material. The text and line-drawings may be reproduced xerographically, 
but glossy prints at publication scale must be supplied for all half-tone illustrations 
and photographic plates. These prints should be identified clearly on the back. 


Referenced publication titles must be spelled out in their entirety. Refer to 
a recent number in the series for details of reference format. Citations of illus- 
trations within the monograph bear initial capitals (e.g., Plate, Text-figure), but 
citations of illustrations in other articles appear in lower-case letters (e.g., plate, 
text-figure). 


Original plate photomounts should have oversize cardboard backing and 
strong tracing paper overlays. These photomounts should be retained by the author 
until the manuscript has been formally accepted for publication. Explanations of 
text-figures should be interleaved on separate numbered pages within the text, 
and the approximate position of the text-figure in the text should be indicated. 
Explanations of plates follow the References Cited. 


Submissions are welcome from any author, regardless of institutional or 
organizational affiliation. Authors must, however, be members of the Paleonto- 
logical Research Institution at the time of publication; annual membership is 
currently US$20.00. Authors are currently required to pay illustration charges. 
Contact the editor for details. 


Collinson, J. 
1962. Size of lettering for text-figures. Journal of Paleontology, vol. 36, p. 
1402. 


Gilbert Dennison Harris 
(1864 - 1952) 


Founder of Palaeontographica Americana (1916) 


ISBN 0-87710-435-7